BanningAnthonysamy Whitney

SPATIAL ECOLOGY, HABITAT USE, GENETIC DIVERSITY, AND
REPRODUCTIVE SUCCESS: MEASURES OF CONNECTIVITY OF A

SYMPATRIC FRESHWATER TURTLE ASSEMBLAGE IN A
FRAGMENTED LANDSCAPE
BY
WHITNEY JOANNA BANNING ANTHONYSAMY
DISSERTATION
Submitted in partial fulfillment of the requirements
for the degree of Doctor of Philosophy in Natural Resources and Environmental Sciences
in the Graduate College of the
University of Illinois at Urbana-Champaign, 2012
Urbana, Illinois
Doctoral Committee:
Professor Jeffrey D. Brawn, Chair
Affiliate Professor Christopher A. Phillips, Director of Research
Affiliate Professor Marlis R. Douglas
Assistant Professor Robert L. Schooley
Associate Professor Carla E. Cceres
ABSTRACT
Habitat fragmentation can have serious conservation implications for long-lived species
such as freshwater turtles. Using integrative radio-telemetry and molecular methods, I examined
characteristics in five species of turtles that should influence connectivity and long-term
persistence of populations among remnant preserves within the Lower Des Plaines River Valley,
a fragmented landscape in northeastern Illinois. Comparisons of movement and habitat use
among Blandings turtle (Emydoidea blandingii), spotted turtle (Clemmys guttata), painted turtle
(Chrysemys picta), common snapping turtle (Chelydra serpentina), and eastern musk turtle
(Sternotherus odoratus) revealed that E. blandingii made long distance movements and readily
moved between wetlands, whereas the other species were more restricted to aquatic movements.
However, S. odoratus, C. serpentina, and C. picta were also capable of making long distance
aquatic movements ( 1 km) via the Des Plaines River. Conversely, C. guttata exhibited the
shortest movements and smallest home range. Patterns of macro- and micro-habitat use
demonstrated strong partitioning between C. guttata and C. picta, C. serpentina, S. odoratus as
well as broad measures of niche breadth and niche overlap for E. blandingii and C. serpentina.
These results suggest that E. blandingii and C. serpentina are habitat generalists whereas C.
guttata is a habitat specialist. Differences in movement and habitat use were likely caused by
species-specific traits and requirements and can impact levels of gene flow within species in
fragmented landscapes. Using microsatellite DNA markers, I examined population genetic
structure in E. blandingii, C. picta, and C. serpentina. I observed moderate to high levels of
genetic diversity in all three species. I detected significant pairwise FST divergence in E.
blandingii between an intact site and three fragmented sites as well as between two fragmented
sites and in C. serpentina between two fragmented sites. Gene flow was male-biased in E.
blandingii across the fragmented sites but differences in patterns of dispersal between males and
ii
females in C. picta and C. serpentina were weak. I found no evidence of genetic population
bottlenecks in any species, but simulations of future genetic diversity suggest that E. blandingii
is more vulnerable to loss of genetic diversity than C. picta or C. serpentina. Finally, I evaluated
the mating system of E. blandingii by corroborating field observations of mating attempts during
radio-telemetry surveys with genetic parentage analysis. I observed promiscuous mating
behavior in E. blandingii as males and females engaged in mounting behaviors with multiple
individuals. Males and females mated successfully with multiple individuals, but successful
matings did not always correspond with observed mating attempts and parentage was strongly
skewed in males. For males, the number of successful mates was positively correlated with total
number of offspring sired. Correlation between relatedness of male-female pairs and
reproductive success was not evident. Repeat paternity in clutches among years was common but
I only documented one confirmed instance of across-season sperm storage. I also only detected
8% multiple paternity in 28 clutches. High variation in reproductive success and low levels of
multiple paternity may be attributed to small population size. During this study, I detected
differences among species in traits such as vagility, niche breadth, and future levels of genetic
diversity. These differences are likely related to species-specific life history traits and should
differentially influence how each of these species responds to fragmentation.
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ACKNOWLEDGEMENTS
It has truly been a privilege to work on this turtle project and I have many people and
organizations to thank for giving me this invaluable opportunity. I am grateful to Dr. Chris
Phillips for inviting me to join his lab eight years ago and the for patience and support he has
afforded me throughout my graduate career, Dr. Marlis Douglas for her teaching and advice on
genetics as well as her mentorship, and Dr. Bob Schooley, Dr. Jeff Brawn, and Dr. Carla Cceres
for imparting their instrumental expertise and guidance throughout the process of completing my
dissertation. This opportunity would also not have been possible without the collaboration,
contribution, and friendship of Dr. Mike Dreslik, Dave Mauger, Natalie Marioni, Dan
Thompson, and Dan Kirk as well as the funding and support provided by the Illinois Toll
Highway Authority, Forest Preserve District of Will County, Forest Preserve District of Dupage
County, Illinois Department of Natural Resources, Chicago Wilderness, Chicago Herpetological
Society, Illinois Academy of Sciences, University of Illinois Urbana Champaign, the Prairie
Research Institute, and the Illinois Natural History Survey. I am indebted to all of the field
assistants who worked so diligently, even in unpleasant environmental conditions, to help collect
the data for my dissertation; Lauren Noffke, Carl Schmidt, Cassandra Sung, Sarabeth Klueh,
Peter Markos, Rachel Bradfield, Christina Aiello, Jeanne Baker, Mike Mosher, Jess Stephens,
Laura Pratt, Laura Lewis, Tyler Pedersen, Mike Knoerr, Linda Rusak, Erin Wilichowski, Ben
von Korf, Teal Richards Dimitrie, Jennifer Heeymeyer, Susan Dalgarn, and Luke Hodges. I
especially thank Jason Ross for his dedication and overall contribution to the turtle project. I
thank Paul Tinnerella for his guidance in the molecular lab and teaching me how to perform
essential lab protocols. For their dependable and meticulous assistance in the lab, I thank Brian
Clague and Stacy Beyer. My graduate experience would not have been as enjoyable and
productive without the friendships and contributions of my colleagues in the herpetology lab;
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John Petzing, Jen Mui, Anne Readel, Andrew Kuhns, Evan Menzel, Jon Warner, Chris Benda,
Brad Cosentino, Sarah Wylie, Dan Wylie, Abby Berkey, Andrew Berger, Ellen Schneider, and
Tanya Hawley. My sincere gratitude also extends to my wonderful friends Michelle and Dan
Neuhauser, Erik and Kim Oslawski, Whitney Cox, and Marilyn Strl. Finally, I thank my family,
especially my parents, Christy and Randy Banning for their perpetual support and
encouragement in my decision to pursue my passion in wildlife ecology and conservation, my
husband Allan for his endless support and uplifting humor, my grandma Margaret Banning, my
sister Shannon and her husband Brad Wilson, and my sister and brother-in law Adal and Matt
Ungerank for all their support and kindness.
TABLE OF CONTENTS
CHAPTER 1: Spatial ecology of a freshwater turtle assemblage in a fragmented landscape .....1

Literature Cited ............................................................................................................................26
Tables ...........................................................................................................................................33
Figures..........................................................................................................................................39
CHAPTER 2: Habitat partitioning in five sympatric freshwater turtle species at an isolated
preserve. .......................................................................................................................................51
Tables ...........................................................................................................................................74
Figures..........................................................................................................................................78
CHAPTER 3: Comparison of population genetic structure among three sympatric freshwater
turtle species.................................................................................................................................84
Tables ...........................................................................................................................................116
Figures..........................................................................................................................................122
CHAPTER 4: Mating system and reproductive success in a fragmented population of Blandings
turtles (Emydoidea blandingii) ....................................................................................................126
Tables ...........................................................................................................................................161
Figures..........................................................................................................................................170
CHAPTER 5: Summary ...............................................................................................................174
APPENDIX A: Spatial metrics for Emydoidea blandingii ..........................................................184
APPENDIX B: Spatial metrics for Clemmys guttata ..................................................................187
APPENDIX C: Spatial metrics for Sternotherus odoratus ..........................................................189
APPENDIX D: Spatial metrics for Chelydra serpentina ............................................................190
APPENDIX E: Spatial metrics for Chrysemys picta ...................................................................191
APPENDIX F: Sample sizes for habitat partitioning analyses ....................................................192
APPENDIX G: Habitat partitioning post-hoc statistical results ..................................................193
APPENDIX H: Multiplex panels .................................................................................................194
APPENDIX I: Genetic Diversity Indices ....................................................................................195
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APPENDIX J: Number of potential and successful mates ..........................................................200
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CHAPTER 1
SPATIAL ECOLOGY OF A FRESHWATER
TURTLE ASSEMBLAGE IN A FRAGMENTED LANDSCAPE
INTRODUCTION
Understanding the consequences of habitat fragmentation requires knowledge about an
organisms life history and ecological traits (e.g. reproductive effort, generation time, body size,
dispersal ability, habitat specialization; Henle et al., 2004; Ewers and Didham, 2006). Sensitivity
to fragmentation depends on a species vagility, the ability to move through a landscape, with
less mobile species often suffering more negative effects than more mobile species in fragmented
landscapes (Lens et al., 2002; ckinger et al., 2009; ckinger et al., 2010). Thus, this chapter
will focus on vagility as a determinant of species responses to fragmentation.
Reptiles have restricted mobility compared to most other vertebrate taxa. Although some
species of freshwater turtles are known to move several kilometers during nesting forays and
among wetlands (Ernst and Lovich, 2009), such movements are often prevented by
anthropogenic barriers such as roads and railroad tracks (Aresco, 2005; Kornilev et al., 2006;
Shepard et al., 2008). Many turtle species are imperiled because of fragmentation (Mitchell and
Klemens, 2000), and persistence of species depends on the ability of individuals to move both
within populations (e.g. among habitat types) and among populations (immigration-emigration
processes).
Using radio-telemetry methods, I examined the spatial ecology for two locally rare and
three common sympatric turtle species occurring in a fragmented landscape. My objectives were
to 1) compile home range and movement parameters for each species, 2) test for differences in
these spatial metrics among sex and stage class within species, 3) test for differences in spatial
1
metrics among populations within species, 4) test for correlation between body size and home
range size within each species, and 5) test for differences in spatial metrics among species.
Specifically, I was interested in addressing the following questions about spatial ecology within
and among species: 1) Does home range and movement differ among stage/sex classes within
each species? 2) Do these spatial metrics differ among sites within species? 3) Does body size
within stage/sex groups influence home range size? and 4) Does home range and movement
differ among species?
I expected that spatial metrics would differ among stage/sex class within species because
life history strategies vary between adults and juveniles between males and females. For
example, life history strategies of juvenile E. blandingii are concentrated on growth and
overcoming low survival rates (Congdon et al., 1993), and most activity appears to be limited to
specific habitat areas that provide better foraging opportunities and refugia (Pappas and Brecke,
1992). Thus, I predicted adults to have larger movements and home range areas than juvenile E.
blandingii. Further, because differences in reproductive strategies (mate searching vs. nesting)
between males and females are predicted to influence movement and activity (Morreale et al.,
1984), with the exception of E. blandingii, I predicted that males in the remaining species to
have larger movements and home range areas than females. Both male and female E. blandingii
are known to make long-distance movements (Rowe and Moll, 1991, Sexton, 1995; Piepgras and
Lang, 2000; Joyal et al., 2001), thus I predicted no differences in spatial metrics between males
and females in E. blandingii. In addition, because E. blandingii are considerably vagile (Rowe
and Moll, 1991, Sexton, 1995; Piepgras and Lang, 2000; Joyal et al., 2001), use multiple habitat
types, and have a larger estimate of niche breadth (Chapter Two), I also expected E. blandingii to
have larger estimates of home range size and movement than C. guttata, S. odoratus, C. picta,
and C. serpentina.
METHODS
Study Site
The Lower Des Plaines river valley (LDPRV) was once a prairie-dominated landscape
(Bowles and McBride, 2001) composed of semi-contiguous, prairie-wetland matrices that would
have allowed turtles to disperse along the river corridor without anthropogenic impediment.
Since the early 1800s there have been drastic changes (e.g. agriculture, roadways, industrial
parks, quarries, shipping canals) to the LDPRV landscape. Remaining natural areas are
effectively isolated from one another except for their connection along the usually very narrow
Des Plaines River and its riparian zone. This study took place at three of these remnant areas in
Will County, Illinois; Will 1 (95 ha), Will 2 (188 ha), and Will 3 (124 ha). Each of these
preserves was inhabited by an abundant turtle fauna including state-listed species such as the
Blandings turtle (Emydoidea blandingii) and spotted turtle (Clemmys guttata) as well as three
common species, the common snapping turtle (Chelydra serpentina), painted turtle (Chrysemys
picta), and common musk turtle (Sternotherus odoratus).
Radio-telemetry
Selected numbers of turtles were radio-tagged and tracked for varying lengths of time
(depending on species and location) during 2005-2010. Radio-tagged turtles were located during
at least two months of one active season (April-October). I radio-located C. serpentina, C. picta,
and S. odoratus at the Will 3 site, C. guttata at the Will 2 and Will 3 sites, and E. blandingii at
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Will 1, Will 2, and Will 3 sites. I outfitted transmitters (Holohil Systems Ltd., Carp, ON,
Canada; Wildlife Materials International Inc., Murphysboro, IL, USA; and L.L. Electronics,
Mahomet, IL, USA) to the right or left posterior portion of the carapace. For individuals < 175 g,
I adhered transmitters by gently abrading the shell with sand paper, applying a small amount of
quick drying epoxy (Marine Power PC11) around the transmitter, and then molding it firmly to
the shell with masking tape (which was removed after the epoxy dried). For individuals >175 g, I
used either the epoxy method or I drilled 1-2 holes in the marginal scutes and securely bolted a
transmitter package constructed of aluminum flashing, plasti-dip, and epoxy. Transmitter
package weight did not exceed 10% of the individuals mass. Stage/sex class (adult male, adult
female, and juvenile) was assigned based on the presence or absence of secondary sexual
characteristics (e.g. concavity of plastron, elongated foreclaws, position of cloaca relative to the
posterior edge of carapace) and sizes of maturation based on previous studies (Ernst and Lovich,
2009). I tracked turtles approximately 3-7 times per week during the active season (April
October) and reduced the frequency of locations to1-2 times per month during over-wintering
(November-March). At each location, I recorded GPS coordinates (UTM-NAD 83 CONUS).
Estimation of spatial parameters

I plotted all turtle location coordinates on an aerial photograph of the preserves using
ArcView 3.2. I included nesting movements of gravid females because these movements and
locations represent areas critical for reproduction. Using the Spatial Analyst and Animal
Movement extensions, I generated movement paths, location statistics, and home ranges for each
individual (Hooge and Eichenlaub, 1997). I calculated mean daily distance (MDD) using only
locations collected one or two days apart from each other during the active season to reduce
under-estimation of actual movements
I estimated home range size in ArcView using multiple methods: minimum convex
polygon (MCP), home range length (HRL), 95% fixed kernel density isopleths (95K), and 50%
fixed kernel density isopleths (50K). I counted the number of 50% isopleth activity centers (core
activity centers, #C) for each individual. Multiple methods were used so that comparisons could
be made easily to other studies. In addition, providing different estimates of home range methods
alleviate criticisms associated with approaches. For example, multiple convex polygons (MCP)
tend to over-estimate home range use by including areas not used by an individual and size is
often correlated with number of locations (Worton, 1987). Because kernel density estimates are a
function of the time an organism spends in an area, they are often better predictors of actual area
use than MCP (Worton, 1987; Seaman and Powell, 1996). However, kernel density estimates can
exclude important areas that are infrequently used, such as overland corridors among wetlands or
critical habitats (i.e. nesting areas) that are important for conservation planning. Home range
length (HRL) was measured as the distance between the two farthest locations and used to
indicate how far an individual was able to transverse during the study; this information was not
always conveyed by home range area estimates. Number (#C) and size of core area use (50K)
was used to evaluate differences in routine area use (e.g. daily foraging).
Frequent radio-locations can lead to non-independence among locations within
individuals. However, autocorrelation has little effect on accuracy of kernel density estimates
and subsampling locations to reduce autocorrelation decreases sample size and accuracy of home
range estimates (De Solla et al., 1999). Thus, all radio-locations were included when estimating
home range parameters.
For kernel estimates, I calculated the smoothing factor (h-values) by averaging the ad hoc
default generated via least squares cross validation (LSCV) for each turtle over the study
duration (Seaman and Powell, 1996). I constructed area curves by plotting MCP size of
sequential samples and MCP size of random samples. I generated sequential MCP areas in
Biotas 1.03a (Ecological Software Solutions LLC) and random MCP areas from bootstrapping
(100 samples) using the Animal Movement Extension. I determined that a sufficient number of
locations had been obtained to represent home range for each turtle when area curve plots were
asymptotic.
Within-species comparisons
Because, E. blandingii and C. guttata were monitored across multiple sites and multiple
stage/sex classes, I used a two-way ANOVA to compare MCP, 95K, 50K, MDD, and HRL
among sites, between stage/sex class, and stage/sex class*site. For significant effects within E.
blandingii, I followed the ANOVA with Gabriels multiple comparison for unequal sample sizes.
Because S. odoratus, C. serpentina, and C. picta were only tracked at one site, I used a Students
t-test to test for differences in all spatial variables only between stage/sex class. Number of core
areas (#C) did not meet the assumptions of normality, thus, I used non-parametric tests to
compare #C among and between sites and stage/sex class. For E. blandingii, I used KruskalWallis tests to compare #C by stage/sex class and by site. For C. guttata, S. odoratus, C.
serpentina, and C. picta, I used a Mann-Whitney U-test to compare #C by stage/sex class and by
site when applicable. Finally, I tested for correlations between carapace length (CL) and home
range (MCP and 95K) to determine if body size influences home range size.
Among-species comparisons
To compare home range and movement among species, I pooled males and females from
the Will 3 site and recalculated MCP, MDD, and HRL to include only locations collected from
May-September 2006. I excluded 2005 data to control for between-year variation because 2005
was a drought year (Anthonysamy et al. in review) and most of my telemetry subjects were E.
blandingii in 2005. In addition, I excluded E. blandingii and C. guttata from the Will 1 and Will
2 sites from this analysis because not all species were tracked at all sites and site-effect
differences could bias results. Because I sampled only adult individuals in the other species, I
also excluded juvenile E. blandingii from this analysis. Kernel estimates (95K, 50K, and #C)
were not used in this analysis because smoothing factors generated for kernel estimates were
species-specific and invalidated statistical comparisons among species. I used a one-way
ANOVA followed with a Gabriels multiple comparison for unequal sample sizes to compare
MCP, MDD, and HRL among species.
For all statistical tests, I tested the assumptions of normality and homogeneity of
variables using the Shapiro-Wilk test and Levenes test, respectively. Variables were Log10+1
(MCP, 95K, 50K), Log10 (HRL), or Ln (MDD) transformed for parametric tests when necessary
to meet the assumptions. I conducted statistical analyses in SPSS 17.0 (SPSS Inc. Chicago,
Illinois) and accepted significance at the 95% level except for post hoc comparisons.
Significance levels for post hoc tests were adjusted with Bonferroni correction and are reported
in the results. Home range and movement parameters are reported as mean 1 S.E.
RESULTS
Within-species comparisons rare species
Emydoidea blandingii
I was unable to obtain a sufficient number of locations for area curves to asymptote in 11
individuals. For the remaining 69 E. blandingii, I collected 7210 locations (277.3 23.8) for
seven males, 15 females, and four juveniles at the Will 1 site from 2006-2009; 3013 locations
(215.2 33.4) for three males, six females, and five juveniles at the Will 2 site from 2007-2009;
and 5390 locations (185.9 13.4) for five males, 14 females, and ten juveniles at the Will 3 site
from 2005-2007 (Appendix A). Turtles were assigned as residents to the site of their original
capture. During this study, two resident turtles (one male and one female) from the Will 2 site
moved to the Will 1 site and back and one resident male from Will 1 moved to the Will 2 site
and back.
Mean values for home range and movement parameters by site and stage/sex class are
shown in Table 1.1 and illustrated Fig(s) 1.1A-E. For males at all sites, minimum convex
polygon home range estimates (MCP) averaged 48.1 10.0 ha, 95% fixed kernel home range
estimates (95K) averaged 14.4 1.4 ha, 50% fixed kernel density isopleths (50K) averaged 1.5
0.1 ha, mean daily distance (MDD) averaged 47.8 5.6 m, and home range length (HRL)
averaged 1507.4 240.5 m. For females at all sites, MCP averaged 26.6 2.8 ha, 95K averaged
13.3 1.0 ha, 50K averaged 1.8 0.2 ha, MDD averaged 34.5 2.7 m, and HRL averaged
1087.9 82.4 m. For juveniles at all sites, minimum convex polygon home range estimates
(MCP) averaged 11.8 3.3 ha, 95K averaged 7.1 0.7 ha, 50K averaged 1.4 0.1 ha, MDD
averaged 20.8 2.2 m, and HRL averaged 721.1 111.6 m.
Two-way ANOVA results are provided in Table 1.2. Minimum convex polygons (MCP)
varied among stage/sex class (F2,60 = 11.52, P 0.0001) and site (F2,60 = 5.3, P = 0.008) but not
by the stage/sex*site interaction term. Post-hoc comparisons (adjusted = 0.0167) revealed that
adult females had larger MCP estimates than juveniles (P 0.0001) and adult males had larger
MCP estimates than juveniles (P 0.0001). No difference in MCP was detected between males
and females. Among sites, turtles at the Will 1 and Will 2 sites had larger MCP estimates than
those at the Will 3 site before Bonferroni correction (P = 0.003 and P = 0.029, respectively). No
difference in MCP was detected between the Will 1 and Will 2 sites.
Ninety-five percent fixed kernel density isopleths (95K) varied among stage/sex class
(F2,60 = 11.38, P 0.0001) but not site. The stage/sex class*site interaction term was not
significant. Post-hoc comparisons revealed that adult females had larger 95K estimates than
juveniles (P 0.0001) and adult males had larger 95K estimates than juveniles (P 0.0001). No
difference in 95K was detected between adult males and adult females. Fifty percent fixed kernel
density isopleths (50K) did not differ among stage/sex class or site.
Mean daily distance (MDD) varied among stage/sex class (F2,60 = 6.60, P = 0.003) and
site (F2,60 = 15.12, P 0.0001) but the stage/sex class*site interaction term was not significant.
Post-hoc comparisons revealed that adult females had significantly greater MDD than juveniles
(P = 0.003) and adult males had greater MDD than juveniles (P 0.0001). No difference in
MDD was detected between adult males and adult females. Among sites, turtles at the Will 1 and
Will 2 sites had greater MDD than those at the Will 3 site (P 0.0001, P = 0.002, respectively).
No difference in MDD was detected between Will 1 and Will 2 sites.
Home range length (HRL) varied among stage/sex class (F2,60 = 5.96, P = 0.004) and site
(F2,60 = 6.04, P = 0.004) but the stage/sex class*site interaction term was not significant. Post-
hoc comparisons revealed that adult females had greater HRL than juveniles (P = 0.004) and that
adult males had greater HRL than juveniles (P 0.0001). No difference in HRL was detected
between adult males and adult females. Among sites, turtles at the Will 1 site had greater HRL
than those at the Will 3 site (P = 0.001). No difference in HRL was detected between Will 1 and
Will 2 or Will 2 and Will 3.
The number of core activity centers (#C) averaged 1.4 0.2 m, 1.4 0.1 m, and 1.2 0.1
m for males, females, and juveniles, respectively. There was no difference among #C for
sex/stage class (2 = 2.577, df = 2, P = 0.276) or site (2 = 5.817, df = 2, P = 0.055). Carapace
length was positively correlated with MCP (r2= 0.547, P 0.0001) and with 95K (r2= 0.566, P
0.0001). Within sex/stage class, carapace length was only correlated with 95K (r2= 0.606, P =
0.013).
Clemmys guttata
I was unable to obtain a sufficient number of locations for area curves to asymptote in
two individuals. For the remaining 34 C. guttata, I collected 1186 locations (mean = 107.8
16.2) for six males and five females at the Will 3 site during 2005-2006, and 3729 locations
(mean = 162.1 17.3) for 12 males and 11 females at the Will 2 site from 2007-2008 (Appendix
B).
shown in Table 1.3 and illustrated in Fig(s) 1.2A-E. For males at both sites, minimum convex
polygon home range estimates (MCP) averaged 2.2 0.5 ha, 95% fixed kernel home range
estimates (95K) averaged 1.2 0.1 ha, 50% fixed kernel home range estimates (50K) averaged
0.2 0.02 ha, mean daily distance (MDD) averaged 12.2 1.3 m, and home range length (HRL)
10
averaged 261.7 37.7 m. For females at both sites, MCP averaged 3.0 0.8 ha, 95K averaged
1.3 0.1 ha, 50K averaged 0.2 0.0 ha, MDD averaged 14.7 1.7 m, and HRL averaged 328.8
57.9 m.
Two-way ANOVA results are provided in Table 1.2. Minimum convex polygons (MCP)
did not vary among stage/sex class or site. Ninety-five percent fixed kernel density isopleths
(95K) and 50K varied between sites (F1,30 = 7.85, P = 0.009; F1,30 = 6.13, P = 0.019,
respectively) but not between stage/sex class or for the stage/sex*site interaction terms. Mean
daily distance (MDD) was greater for females than males (F1,30 = 40.27, P 0.0001) and greater
in the Will 2 than Will 3 site (F1,30 = 40.27, P 0.0001) but not for the stage/sex*site interaction
term. Home range length (HRL) did not differ among stage/sex class or site.
The number of core activity centers (#C) averaged 1.4 0.2 m and 2.0 0.2 m for males
and females, respectively. Females had a significantly greater #C than males (U = 74.0, P =
0.008) but there were no differences between the Will 2 and Will 3 sites (U = 94.0, P = 0.191). I
found a nearly significant correlation between carapace length and MCP (r2 = 0.332, P = 0.055).
Within stage/sex class, I found a significant correlation between carapace length and MCP (r2=
0.583, P = 0.018) and a nearly significant correlation between carapace length and 95K (r2=
0.484, P = 0.058) only in females.
Within-species comparisons common species

Sternotherus odoratus
three individuals and thus excluded them from analyses. For the remaining 12 S. odoratus, I
11
collected 708 (mean = 59.0 5.0) locations for six males and six females at the Will 3 site from
2005-2006 (Appendix C).
shown in Table 1.4 and illustrated in Fig(s) 1.3A-E. For males, minimum convex polygon home
range estimates (MCP) averaged 11.6 9.3 ha, 95% fixed kernel home range estimates (95K)
averaged 5.0 0.8 ha, 50% fixed kernel home range estimates (50K) averaged 0.9 0.1 ha,
mean daily distance (MDD) averaged 36.3 11.6 m, and home range length (HRL) averaged
585.4 278.1 m. For females, MCP averaged 8.2 4.7 ha, 95K averaged 5.3 1.2 ha, 50K
averaged 1.0 0.2 ha, MDD averaged 30.0 5.6 m, and HRL averaged 589.8 222.3 m. No
difference in MCP, 95K, 50K, MDD, or HRL was detected between males and females (Table
1.5).
and females, respectively. There was no statistically significant difference between #C for
stage/sex class (U = 15.0, P = 0.523). I found no correlation between carapace length and home
range size estimates.
Chelydra serpentina
two individuals and thus excluded them from analyses. For the remaining nine C. serpentina, I
collected 597 locations (mean = 66.3 6.8) for five males and four females at the Will 3 site in
2006 (Appendix D).
12
1.5).
and females, respectively. There was no statistically significant difference between #C for
stage/sex class (U = 7.5, P = 0.264). I found no correlation between carapace length and home
range size estimates.
Chrysemys picta
one individual and thus excluded her from analyses. For the remaining eight C. picta, I collected
379 locations (mean = 47.4 6.6) for five males and three females at the Will 3 site in 2006
(Appendix E).
13
1.5).
The number of core activity centers (#C) averaged 1.0 0.0 m for both, males and
females. There was no statistically significant difference between #C for stage/sex class (U = 7.5,
P = 1.00). I found no correlation between carapace length and home range size estimates.
Among-species comparison
A total of 17 E. blandingii, ten C. guttata, nine S. odoratus, nine C. serpentina, and eight
C. picta were included in the among species comparison. Mean MCP for E. blandingii, C.
guttata, S. odoratus, C. serpentina, and C. picta was 8.8 2.0 ha, 1.6 0.6 ha, 3.2 0.9 ha, 5.8
1.5 ha, and 6.2 1.9 ha, respectively. Mean MDD for E. blandingii, C. guttata, S. odoratus, C.
serpentina, and C. picta was 39.0 5.1 m, 9.0 1.9 m, 25.9 1.8 m, 35.5 7.5 m, and 53.2
22.4 m, respectively. Mean HRL for E. blandingii, C. guttata, S. odoratus, C. serpentina, and C.
picta was 545.2 81.4 m, 238.1 60.1 m, 360.7 52.5 m, 528.9 110.4 m, and 700.5 213.4
m, respectively.
Significant differences in MCP, MDD, and HRL were detected among species of adult
individuals at the Will 3 site (F4,48 = 3.951, P = 0.008; F4,48 = 11.139, P 0.0001; F4,48 =
3.606, P = 0.012, respectively). Post-hoc comparisons (adjusted = 0.0125) revealed that E.
blandingii had significantly greater MCP estimates than C. guttata (P = 0.005; Fig. 1.4A). No
difference in MCP was detected between E. blandingii and the remaining species. Emydoidea
blandingii, S. odoratus, C. serpentina, and C. picta had significantly greater MDD estimates than
C. guttata (P 0.001; Fig. 1.4B). No differences in MDD comparisons were detected between
14
the other species. Emydoidea blandingii and C. picta had significantly greater HRL than C.
guttata before but not after Bonferroni correction (P = 0.019 and P = 0.021; Fig. 1.4C). No
differences in HRL comparisons were detected between the other species.
DISCUSSION
Within-species comparisons rare species
Many radio-telemetry studies report on the spatial ecology of E. blandingii and C. guttata
because of the elevated conservation status of these species throughout their ranges (Ernst and
Lovich, 2009). However, my radio-telemetry studies of E. blandingii and C. guttata in a
fragmented landscape provide extensive data sets with robust estimates of the movement and
home range of these two species. For example, I collected numerous radio-locations for several
individuals of different stage/sex classes during periods of 1 active season across multiple
sites. In comparison, many other studies located far fewer turtles and located individuals less
frequently or over a shorter time period, precluding their ability to estimate a rigorous home
range size for some individuals (Ernst, 1970; McNeil, 2002), statistically compare stage/sex
classes (Graham, 1995; Rubin et al., 2001; Innes et al., 2008), or test independent data (i.e.
pooling multiple observations for single individuals; Ross and Anderson, 1990; Rowe and Moll,
1991). My data sets can be used to establish a firm foundation of spatial ecology on which to
further develop ideas and hypotheses about additional issues of turtle spatial ecology (e.g.
connectivity in a fragmented landscape).
15
Adult E. blandingii populations within the LDPRV averaged larger 95% fixed kernel
home range estimates, mean daily movement distances, and home range length distances than
juveniles. Piepgras and Lang (2000), also reported smaller juvenile home range sizes compared
to adults but found that females and juveniles travel greater straight-line daily distances than
males. Adult E. blandingii are known to make long (> 1 km) inter-wetland forays (Piepgras and
Lang, 2000; Rowe and Moll, 1991) and when traveling to nesting locations (Sexton, 1995;
Piepgras and Lang, 2000; Joyal et al., 2001). In my study, I observed three adult individuals to
move from their resident site to a different adjacent site and then move back to their resident site.
Conversely, because the life history strategies of juvenile E. blandingii are concentrated on
growth and overcoming low survival rates (Congdon et al., 1993), most activity appears to be
limited to specific habitat areas that provide better foraging opportunities and refugia (Pappas
and Brecke, 1992). The inclusion of post-nesting locations in one radio-telemetry study were
thought to be responsible for larger female movements compared to males (Ross and Anderson,
1990) and reproductive class had an effect on home range size in an Ontario population (Millar
and Blouin-Demers, 2011). Although I included nesting locations in estimates of movement and
home range, I found no difference in these parameters between male and female E. blandingii
within the LDPRV. Similar findings were reported for suburban E. blandingii populations in
Massachusetts (252-1246 ha; Grgurovic and Sievert, 2005), an intact population in Ontario (3400
ha; Edge et al., 2010), and a large but historically disturbed site in Wisconsin (3884 ha; Schuler
and Thiel, 2008). In my study, larger males tended to have larger home range sizes than smaller
males.
16
Effects of site location were also important in this study as individuals from the Will 1
site averaged greater movement and home range length distances than individuals from the Will
3 site. Site resource differences, such as the size, type, and distribution of wetland areas and the
proximity of these areas to the Des Plaines River, likely accounted for some of this variation. In
addition, tracking was conducted at different years among sites and differences in habitat
availability among years, could account for site differences. For example, E. blandingii at a
preserve in Will County moved shorter mean daily distances during a drought year compared to
a wet year (Anthonysamy et al. in review). Core area (50% kernel estimate size and number) use
was similar among all individuals within LDPRV regardless of stage/sex class or site and
primarily represented intra-marsh foraging movements. Many E. blandingii spatial ecology
studies report on the number of activity centers and differences in number of activity centers
among stage/sex classes (Ross and Anderson 1990; Rowe and Moll, 1991; Piepgras and Lang,
2000; Innes et al., 2008) but wide variation in core area (i.e. activity center) definition and
estimation exists among these studies. Thus, it is difficult to make comparisons of core area use
between other studies and LDPRV.
Average MCP areas for adult LDPRV turtles (males = 48.1 ha; females = 26.6 ha) fell
within the range of estimates reported for other studies (Ernst and Lovich, 2009) but were large
compared to seasonal MCP estimates reported for populations in another urban Illinois landscape
of similar size (Rubin et al., 2001) and comparable to MCP estimates for a large Minnesota
population (Piepgras and Lang, 2000). The relatively large MCP estimates for LDPRV turtles
may be a result of multi-year radio-tracking for some individuals. Schuler and Thiel (2008)
showed that E. blandingii home range size increases linearly with monitoring duration over
multiple years. However, comparisons of my LDPRV 95% fixed kernel home range estimates
17
were smaller than kernel estimations for the Ontario and Massachusetts studies (Grgurovic and
Sievert, 2005; Edge et al., 2010) suggesting that kernel estimators may serve as better home
range comparisons among studies than MCP. Only two previous studies documented home range
for juvenile E. blandingii (Piepgras and Lang, 2000; Innes et al., 2008). Average juvenile MCP
and 95K size (11.8 ha and 7.1 ha, respectively) for LDPRV was comparable to Minnesota MCP
size (12.8 ha) and larger than a single juvenile 95% MCP home range size (3.3 ha) in New
Hampshire (Innes et al., 2008).
Clemmys guttata
Female C. guttata populations within the LDPRV averaged greater mean daily movement
distances than males but this did not produce significant differences in home range estimates or
home range length between the sexes. Similarly, no differences in MCP home range were found
between males and females in Pennsylvania (Ernst, 1970) and Ontario (Rasmussen and Litzgus,
2010) or in MCP and home range length in Massachusetts (Milam and Melvin, 2001). However,
differences in MCP home range between males and females were previously detected at the Will
3 site and South Carolina populations when including locations of gravid females (Wilson, 1994;
Litzgus and Mousseau, 2004). Thus, differences in movement and numbers of core home range
areas between male and female C. guttata in the LDPRV is likely attributed to nesting forays of
gravid females. Other studies have reported movement differences among seasons (Litzgus and
Mousseau, 2004; Rasmussen and Litzgus, 2010) but I did not test for seasonal effects in the
LDPRV populations. The positive correlation between body size and home range size in females
may have resulted from the lack of nesting migrations in smaller, immature individuals.
18
The effect of site was also important for C. guttata within LPDRV as individuals from
the Will 2 site averaged greater home range estimates (95K and 50K) and mean daily distance
than individuals from the Will 3 site. As noted above for E. blandingii, differences in resource
distribution between sites or year effects, and tracking duration (one year at Will 3 vs. two years
at Will 2) could have accounted for some of this variation. Core area (50K) was similar between
stage/sex classes but was greater for individuals at the Will 2 than Will 3 site. The number of
core areas used was greater for females than males, and possibly accounts inter-wetland use for
nesting forays, but differences were not evident between sites.
Home range estimates and home range length for C. guttata within the LDPRV fell
within ranges reported in most other studies (Ernst and Lovich, 2009) but were smaller than
those estimated for one study in Ontario (6.5-7.9 ha; Rasmussen and Litzgus, 2010). Besides the
previous study in Will County by Wilson (1994; males = 0.7 ha; females = 1.8 ha; 1994), the
MCP home range size and length of the LDPRV populations most closely resembled those of C.
guttata populations in central Massachusetts (males = 1.9 ha, 261 m; females = 4.6 ha, 345 m;
Milam and Melvin, 2001) and Victoria County, Ontario (males = 3.6 ha; females = 4.7 ha;
Haxton and Berrill, 1999).
Within-species comparisons common species

Although they are more abundant and widely distributed, common species are often less
studied than rare species. Only a few radio-telemetry studies have assessed aspects of spatial
ecology for common species such as C. picta (Rowe, 2003; Rowe and Dalgarn, 2010), S.
odoratus (Rowe et al., 2009), and C. serpentina (Obbard and Brooks, 1980; Obbard and Brooks,
1981; Brown and Brooks, 1993). Previous studies examining home range and movement have
19
also been geographically limited and may not represent the complete range of spatial metrics for
a particular species throughout its distribution. Also, inconsistencies in home range estimates and
movement distances reported among the few studies may result from the use of different field
techniques (trapping vs. radio-telemetry) and not necessarily from variation in spatial metrics
among turtles. For example, many movements go undetected during trapping surveys compared
to radio-telemetry surveys and this disparity makes generalizations between such studies
problematic.
For common species, loss of habitat and increased isolation also has detrimental impacts
including increased road mortality and skewed sex ratios (Aresco, 2005). Further, small
decreases in survival rates of adults are predicted to cause drastic population declines, even in a
common turtle species (Congdon et al., 1994). Yet the impacts of fragmentation on populations
of common turtle species have not been well documented.
Sternotherus odoratus
I detected no differences between male and female S. odoratus for any home range or
movement parameters at the Will 3 site. Rowe et al. (2009), the only other radio-telemetry study
on S. odoratus, also found no differences in home range estimates between sexes. However,
trapping studies documented that male S. odoratus moved longer distances and more frequently
between recaptures than females (Mahmoud, 1969; Ernst, 1986; Smar and Chambers, 2005). In
the present study, small sample sizes of male (N=6) and female (N=6) S. odoratus as well as
individual variation may have prevented the detection of significant differences in home range
and movement estimates between sexes.
20
The 95% fixed kernel density isopleth home range estimate for a population in Michigan
(2.8 ha) was smaller compared to S. odoratus at the Will 3 site (5.1 ha) but 50% fixed kernel
density isopleths (core areas) were similar between the studies, 1.5 ha and 1.0 ha, respectively
(Rowe et al., 2009). Turtles in both studies used 1-2 core areas (Rowe et al., 2009). Average
home range size, estimated from trapping data, for a population in Pennsylvania was also smaller
(males = 1.8 ha; females = 0.9 ha; Ernst, 1986) than in S. odoratus at Will 3 site, but these
estimates were derived from recapture locations and likely underestimated home range size.
In Virginia, S. odoratus displayed site fidelity to ponds suggesting movement was limited
and home ranges were small (Holinka et al., 2003). Yet I documented long-distance movements
of S. odoratus at the Will 3 site as individuals made inter-wetland movements between ponds
and river habitat and completed long forays > 1 km within the Des Plaines River. Other studies,
including displacement studies, have also reported long-distance movements for S. odoratus
(Ernst, 1986; Holinka et al., 2003; Smar and Chambers, 2005; Andres and Chambers, 2006;
Rowe et al., 2009).
Chelydra serpentina
Previous thorough studies on the movement and home range of C. serpentina (Obbard
and Brooks, 1980; Obbard and Brooks, 1981; Brown and Brooks, 1993; Pettit et al., 1995) are
geographically limited (restricted to Ontario, Canada) considering the widespread distribution of
this species in North America. In previous radio telemetry studies, no difference in home range
size was found between males and females at Algonquin Park, Ontario (Obbard and Brooks,
1981) but differences in seasonal movement between the sexes were observed at the same
location (Brown and Brooks, 1993). At Hamilton Harbor, Ontario, female C. serpentina were
21
observed to have larger home ranges and move longer distances than males (Pettit et al., 1995). I
observed marked variation in home range or movement estimates among individuals but failed to
detect differences between male and female C. serpentina at Will 3 site. Average MCP home
range estimates for male C. serpentina in Ontario (3.2 ha) were comparable to males at the Will
3 site (3.9 ha) but estimates for females at the Will 3 site (8.1 ha) were larger than females at
Ontario (3.8 ha; Obbard and Brooks, 1981). Small sample size of males (N=5) and females
(N=4) may have prevented the detection of significant differences in home range and movement
estimates between sexes in my study.
Chelydra serpentina has been reported as being sedentary and inactive (Ernst and Lovich,
2009). However, mean daily distance of C. serpentina in my study averaged 34.5 m and home
range length for two individuals approached 1 km suggesting that this species is moderately
active and capable of long distance movements at the Will 3 site. Radio-telemetered turtles were
typically re-located in the same area for several days at a time but individuals would occasionally
make inter-wetland movements or long forays within the Des Plaines River. I did not assess the
reproductive status nor did I observe nesting of radio-telemetered C. serpentina in my study but
nesting females are capable of moving multiple kilometers over a few days (Obbard and Brooks
1980; Pettit et al., 1995). Reports of inactivity in C. serpentina could be a result of the
misclassification of inactive turtles (e.g. inactive turtle moved when approached and vice versa)
or a bias in the ability to observe active turtles versus inactive turtles (Obbard and Brooks, 1981).
Chrysemys picta
I failed to detect significant differences in home range or movement parameters between
males (N=5) and females (N=3) but this could be attributed to small sample size. However, in
22
previous radio telemetry studies, no differences were observed in home range or movement
parameters among male, female, and juvenile C. picta in Michigan (Rowe, 2003; Rowe and
Dalgarn, 2010). The average mean daily distance (MDD) of 47.4 m/day in my study was shorter
than estimates (68.1- 96.5 m/day) for C. picta in Michigan (Rowe, 2003; Rowe and Dalgarn,
2010). The turtles in my study were radio-located less frequently (once per day) than the
Michigan studies (three times per day) which likely underestimated total daily movement and
accounted for the shorter movement distances in the Will 3 site turtles. However, average MCP
home range estimates for C. picta in Michigan (males = 2.9 ha; females = 1.8 ha) (Rowe and
Dalgarn, 2010) were smaller in comparison to estimations for turtles at the Will 3 site (males =
7.5 ha; females = 3.9 ha). This could be because the C. picta in my study were radio-located at a
wetland complex consisting of marsh, pond, and river habitats whereas the Michigan study
occurred at a small marsh system (Rowe, 2003; Rowe and Dalgarn, 2010).
Considering the widespread abundance of C. picta, few other studies have examined the
spatial ecology for this common species (Pearse, 1923; Sexton, 1959; Gibbons, 1968; McAuliffe,
1978; MacCulloch and Secoy 1983; House et al. 2010). Reported movement distances vary
widely and are dependent on the type of habitat system where the turtles are studied. For
example, distances transversed by C. picta bellii from a river system in Saskatchewan during
trapping studies (MacCulloch and Secoy, 1983) were greater than distances reported for the same
sub-species at a pond complex in a trapping study conducted in Kansas (House et al., 2010).
Additionally, variation in movement among individuals at the Will 3 site tended to correspond
with habitat use. For example, individuals that used the Des Plaines River traveled longer
distances and had larger home range estimates than individuals solely occupying marsh or pond
habitats. Inconsistencies in reported movement distances are also likely a result of the use of
23
different field techniques (trapping vs. radio-telemetry). As stated above, many movements go
undetected during trapping surveys compared to radio-telemetry surveys and this disparity makes
comparisons between the few studies problematic.
Among-species comparison
Common and rare reptile species demonstrate different sensitivities to fragmentation
(Attum et al., 2008). Although turtles are classified as long-lived organisms with low juvenile
recruitment and high adult survival, variation in life history and ecology traits (i.e. ecological
tolerance, vagility, generation time, clutch size, diet, etc.) exists among turtle species (Ernst and
Lovich, 2009) that should impact how they respond to fragmentation. For example, generalist
(common) species are suggested to be more tolerant of fragmentation than specialist (rare)
species (Henle et al., 2004; Ewers and Didham, 2006).
Species included in my study (E. blandingii, C. guttata, S. odoratus, C. serpentina, and
C. picta) exhibit variation in their vagility and habitat specialization (Chapter Two), and I
expected differences in home range size and movement. Within the LDPRV, E. blandingii had
significantly larger MCP home range estimates than C. guttata. Because they are capable of
making long overland forays between wetlands and to nesting sites, E. blandingii are
considerably vagile (Ernst and Lovich, 2009). The ability to transverse the preserve as well as
use a number of different habitat types (Chapter Two) likely contributed to the larger home range
estimates for this species. However, E. blandingii home range length (HRL) was only
significantly larger than C. guttata, indicating that S. odoratus, C. picta, and C. serpentina are
also capable of making long-distance movements. The primary difference in mobility patterns
between E. blandingii and the common species was that long distance movements by S.
24
odoratus, C. serpentina, and C. picta were mostly restricted to within wetlands (i.e. the Des
Plaines River) whereas E. blandingii moved among wetlands.
Clemmys guttata made smaller daily movement distances compared to all other species.
This is likely because C. guttata at the Will 3 site are restricted to concentrated areas of the
preserve that predominantly consist of shallow, sedge-marsh habitat (Chapter Two). Except for
S. odoratus, C. guttata is also the smallest of the five species and may have lower energy
requirements. The other species typically use deeper and more open-water habitats (i.e. ponds,
river) that are conducive to larger movements (Chapter Two). Failure to detect further
differences in some parameters between species could be attributed to small samples sizes in the
common species.
25
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Schuler, M., and R.P. Thiel. 2008. Annual vs. multiple-year home range sizes of individual
Blandings turtles, Emydoidea blandingii, in Central Wisconsin. Canadian FieldNaturalist 122:61-64.
Seaman, D.E., and R.A. Powell. 1996. An evaluation of the accuracy of kernel density estimators
for home range analysis. Ecology 77:2075-2085.
Sexton, O.J. 1959. Spatial and temporal movements of a population of the painted turtle,
Chrysemys picta marginata (Agassiz). Ecological Monographs 29:113-140.
31
Sexton, O. J. 1995. Miscellaneous comments on the natural history of Blandings turtle

(Emydoidea blandingii). Transactions of the Missouri Academy Science 29:1-13.
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Wilson, T.P. 1994. Ecology of the spotted turtles, Clemmys guttata, at the western range limit.
Masters Thesis, Eastern Illinois University, Charleston, Illinois. 97 pp.
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Modelling 38:277-298.
32
TABLES
33
M
F
J
Will 3
M
F
J
Will 2
M
F
J
Will 1
213.4 6.2
199.5 3.3
125.3 5.8
CL (mm)
203.7 13.5
200.2 3.0
118.0 10.5
CL (mm)
219.0 6.9
205.0 3.1
147.3 6.5
CL (mm)
167.6 25.8
210.5 21.9
160.5 18.1
# Loc
287.3 12.4
216.8 56.7
170.0 63.2
# Loc
276.3 36.2
270.0 35.8
306.5 57.7
# Loc
23.4 6.6
24.9 4.6
7.0 2.7
MCP (ha)
75.8 39.1
33.9 9.6
13.6 3.6
MCP (ha)
56.6 13.6
25.6 3.0
21.3 13.9
MCP (ha)
24.3 5.8
28.8 5.1
15.5 2.3
MDD (m)
50.0 6.9
42.3 6.4
22.3 4.0
MDD (m)
63.4 7.2
37.8 2.7
32.2 2.5
MDD (m)
10.2 1.2
13.2 2.0
6.2 0.8
95K (ha)
13.0 2.2
12.6 1.5
9.1 1.7
95K (ha)
18.2 2.2
13.7 1.2
6.8 1.8
95K (ha)
1.4 0.3
1.9 0.2
1.2 0.1
50K (ha)
1.3 0.2
2.4 0.4
1.8 0.2
50K (ha)
1.6 0.2
1.5 0.2
1.4 0.1
50K (ha)
1.8 0.4
1.7 0.2
1.2 0.1
#C
1.0 0.0
1.0 0.0
1.2 0.2
#C
1.4 0.2
1.3 0.2
1.0 0.0
#C
841.3 168.5
946.8 105.6
568.6 116.4
HRL (m)
1844.3 494.6
1271.2 341.9
720.7 163.9
HRL (m)
1903.8 454.7
1143.8 85.9
1102.9 375.1
HRL (m)
34
Table 1.1 Spatial statistics for 69 E. blandingii radio-tracked at three sites (Will 1-3) in Will County, Illinois from 2005-2010.
Individuals were allocated to three different stage/sex categories; males (M), females (F) and juveniles (J). Listed for each site and
stage/sex are means 1SE for: carapace length (CL), number of radio-locations (#Loc), minimum convex polygon (MCP), mean daily
distance moved (MDD), 95% fixed kernel density isopleth (95K), 50% fixed kernel density isopleth (50K), number of 50% fixed
kernel density isopleths (#C), and home range length (HRL).
Table 1.2 Two-way ANOVA results for comparisons of spatial statistics among stage/sex class
and site for 69 E. Blandingii and 34 C. guttata radio-tracked at three sites (Will 1-3) in Will
County, Illinois from 2005-2010. Listed are: minimum convex polygon home range area (MCP),
95% fixed kernel density isopleth (95K), 50% fixed kernel density isopleth (50K), mean daily
distance moved (MDD), and home range length (HRL) among stage/sex class and site.
.
Variable
Effect
E. blandingii
F
df
p
C. guttata
df
MCP
Stage/sex
Site
Stage/sex * Site
11.520 2,60
5.300 2,60
0.496 4,60
<0.0001
0.008
0.739
1.246
0.101
0.875
1,30
1,30
1,30
0.273
0.752
0.357
95K
Stage/sex
Site
Stage/sex * Site
11.380 2,60
2.281 2,60
1.004 4,60
<0.0001
0.111
0.413
1.233
7.852
2.514
1,30
1,30
1,30
0.276
0.009
0.123
50K
Stage/sex
Site
Stage/sex * Site
2.875 2,60
1.527 2,60
1.551 4,60
0.064
0.225
0.199
1.251
6.129
0.235
1,30
1,30
1,30
0.272
0.019
0.632
MDD
Stage/sex
Site
Stage/sex * Site
6.706 2,60
15.365 2,60
1.106 4,60
0.002
<0.0001
0.362
4.522
40.274
2.446
1,30
1,30
1,30
0.042
<0.0001
0.128
HRL
Stage/sex
Site
Stage/sex * Site
6.219 2,60
6.335 2,60
0.700 4,60
0.004
0.003
0.595
1.102
0.014
0.261
1,30
1,30
1,30
0.302
0.908
0.613
35
M
F
Will 3
M
F
Will 2
110.7 5.5
107.6 1.4
CL (mm)
97.1 2.1
99.5 1.8
CL (mm)
82.7 3.6
138.0 31.6
# Loc
146.6 21.9
179.1 27.3
# Loc
1.7 0.8
3.8 1.8
MCP (ha)
2.5 0.7
2.7 0.8
MCP (ha)
5.8 0.9
9.1 1.3
MDD (m)
15.3 1.1
17.2 1.9
MDD (m)
0.8 0.1
1.1 0.2
95K (ha)
1.4 0.1
1.3 0.1
95K (ha)
0.1 0.0
0.1 0.0
50K (ha)
0.2 0.0
0.2 0.0
50K (ha)
1.0 0.0
2.0 0.3
#C
1.7 0.3
2.0 0.2
#C
246.0 61.6
400.7 156.7
HRL (m)
269.5 49.1
296.0 50.2
HRL (m)
36
Table 1.3 Spatial statistics for 34 C. guttata radio-tracked at two sites (Will 1-2) in Will County, Illinois from 2005-2009. Individuals
were allocated to two different stage/sex categories; males (M) and females (F). Listed for each site and stage/sex are means 1SE
for: mean carapace length (CL), number of radio-locations (#Loc), minimum convex polygon (MCP), mean daily distance moved
(MDD), 95% fixed kernel density isopleth (95K), 50% fixed kernel density isopleth (50K), number of 50% fixed kernel density
isopleths (#C), and home range length (HRL).
# Loc
49.0 6.9
69.0 5.0
# Loc
71.4 7.5
60.0 12.7
# Loc
53.2 8.2
37.7 10.4
CL (mm)
S. odoratus
M
105.7 3.6
F
114.0 2.2
CL (mm)
C. serpentina
M
275.8 12.8
F
253.0 14.0
CL (mm)
C. picta
M
141.0 4.2
F
141.7 8.8
7.5 2.7
3.9 2.1
MCP (ha)
3.9 1.9
8.1 2.1
MCP (ha)
11.6 9.3
8.2 4.7
MCP (ha)
70.8 34.4
24.0 6.2
MDD (m)
28.3 10.8
42.3 10.3
MDD (m)
36.3 11.6
30.0 5.6
MDD (m)
11.1 1.2
7.5 1.4
95K (ha)
2.8 0.9
5.6 1.3
95K (ha)
5.0 0.8
5.3 1.2
95K (ha)
2.3 0.3
1.9 0.1
50K (ha)
0.6 0.1
0.5 0.1
50K (ha)
0.9 0.1
1.1 0.2
50K (ha)
1.0 0.0
1.0 0.0
#C
1.0 0.0
1.3 0.3
#C
1.3 0.2
1.2 0.2
#C
663.3 269.4
762.4 424.2
HRL (m)
434.5 147.2
647.0 169.2
HRL (m)
585.4 278.1
589.8 222.3
HRL (m)
37
Table 1.4 Spatial statistics for 12 S. odoratus, nine C. serpentina, and eight C. picta radio-tracked in Will County, Illinois from 20052006. Individuals were allocated to two different stage/sex categories; males (M) and females (F). Listed for each site and stage/sex
are means 1SE for: mean carapace length (CL), number of radio-locations (#Loc), minimum convex polygon (MCP), mean daily
distance moved (MDD), 95% fixed kernel density isopleth (95K), 50% fixed kernel density isopleth (50K), number of 50% fixed
kernel density isopleths (#C), and home range length (HRL).
Table 1.5 Students t-test results for comparisons of spatial statistics between stage/sex class for
12 S. odoratus, nine C. serpentina, and eight C. picta radio-tracked at Will County, Illinois from
2005-2006. Listed are: minimum convex polygon home range (MCP), 95% fixed kernel density
isopleth (95K), 50% fixed kernel density isopleth (50K), mean daily distance moved (MDD), and
home range length (HRL).
Variable
MCP
95K
50K
MDD
HRL
S. odoratus
t
df
p
-0.075
-0.174
-0.672
0.324
-0.101
10
10
10
10
10
0.942
0.865
0.517
0.753
0.922
C. serpentina
t
df
p
-1.491
-1.998
0.044
-1.099
-0.999
7
7
7
7
7
0.180
0.086
0.966
0.308
0.351
t
1.132
1.982
0.881
1.395
-0.119
C. picta
df
p
6
6
6
6
6
0.780
0.095
0.412
0.213
0.909
38
FIGURES
39
Fig. 1.1 Comparisons of spatial statistics between stage/sex class for 69 E. blandingii radiotracked at three preserves in Will County, Illinois from 2005-2010. Listed are: mean estimates (
1SE) of A) Minimum convex polygon (MCP), B) 95% fixed kernel density isopleth (95K), C)
50% fixed kernel density isopleth (50K), D) mean daily distance moved (MDD), and E) home
range length (HRL).
A)
140
Male
Female
Juvenile
MCP (ha)
120
100
80
60
40
20
0
Will 1
Will 2
Site
Will 3
B)
25
Male
Female
Juvenile
95% KDI (ha)
20
15
10
5
0
Will 1
Will 2
Site
Will 3
40
Fig. 1.1 (cont.)

C)
Male
Female
Juvenile
50%KDI (ha)
2.5
2
1.5
1
0.5
0
Will 1
Will 2
Site
Will 3
D)
80
70
Male
Female
Juvenile
MDD (m)
60
50
40
30
20
10
0
Will 1
Will 2
Site
Will 3
41
Fig. 1.1 (cont.)

E)
2500
Male
Female
Juvenile
HRL (m)
2000
1500
1000
500
0
Will 1
Will 2
Site
Will 3
42
Fig. 1.2 Comparisons of spatial statistics between stage/sex class for 34 C. guttata radio-tracked
at two preserves in Will County, Illinois from 2005-2008. Listed are: mean estimates ( 1SE) of
A) Minimum convex polygon (MCP), B) 95% fixed kernel density isopleth (95K), C) 50% fixed
kernel density isopleth (50K), D) mean daily distance moved (MDD), E) and home range length
(HRL).
A)
Male
Female
MCP (ha)
5
4
3
2
1
0
Will 2
Will 3
Site
95% KDI (ha)
B)
1.8
1.6
1.4
1.2
1
0.8
0.6
0.4
0.2
0
Male
Female
Will 2
Will 3
Site
43
Fig. 1.2 (cont.)

C)
0.3
Male
Female
50% KDI (ha)
0.25
0.2
0.15
0.1
0.05
0
Will 2
Will 3
Site
D)
Male
25
Female
MDD (m)
20
15
10
5
0
Will 2
Will 3
Site
44
Fig. 1.2 (cont.)

E)
Male
600
Female
HRL (m)
500
400
300
200
100
0
Will 2
Will 3
Site
45
Fig. 1.3 Comparisons of spatial statistics between stage/sex class for 12 S. odoratus, nine C.
serpentina, and eight C. picta radio-tracked at a preserve in Will County, Illinois from 20052006. Listed are: mean estimates ( 1SE) of A) Minimum convex polygon (MCP), B) 95% fixed
kernel density isopleth (95K), C) 50% fixed kernel density isopleth (50K), D) mean daily
distance moved (MDD), E) and home range length (HRL).
A)
Male
25
Female
MCP (ha)
20
15
10
5
0
S. odoratus
C. serpentina
Species
C. picta
B)
14
Male
Female
95 % KDI (ha)
12
10
8
6
4
2
0
S. odoratus
C. serpentina
Species
C. picta
46
Fig. 1.3 (cont.)

C)
Male
Female
50 % KDI (ha)
2.5
2
1.5
1
0.5
0
S. odoratus
C. serpentina
Species
C. picta
D)
Male
120
Female
MDD (m)
100
80
60
40
20
0
S. odoratus
C. serpentina
Species
C. picta
47
Fig. 1.3 (cont.)

E)
Male
1400
Female
1200
HRL (m)
1000
800
600
400
200
0
S. odoratus
C. serpentina
Species
C. picta
48
Fig. 1.4 Comparisons of spatial statistics between 17 E. blandingii, ten C. guttata, nine S.
odoratus, nine C. serpentina, and eight C. picta radio-tracked at a preserve in Will County,
Illinois during 2006. Listed are: mean estimates ( 1SE) of A) Minimum convex polygon
(MCP), B) mean daily distance moved (MDD), and C) home range length (HRL).
A)
12
MCP (ha)
10
8
6
4
2
0
Species
MDD (m)
B)
80
70
60
50
40
30
20
10
0
Species
49
Fig. 1.4 (cont.)

C)
1000
HRL (m)
800
600
400
200
0
Species
50
CHAPTER 2
HABITAT PARTITIONING IN FIVE SYMPATRIC FRESHWATER TURTLE SPECIES
AT AN ISOLATED PRESERVE
INTRODUCTION
Resource partitioning is fundamental to community structuring (Schoener, 1974). Empirical
studies demonstrate that species coexist by partitioning resources along multiple gradients such
as food, habitat, time, and space (Luiselli, 2006; Luiselli, 2008; Robertson et al., 2008). Niche
breadth and amount of niche overlap among co-existing species varies depending on phenotypic
and ecological similarities (Pacala and Roughgarden, 1982; Cromsigt and Olff, 2006) as well as
abiotic factors such as the availability of limiting resources (Sebasti, 2004). In a review of
resource partitioning studies in freshwater turtles, habitat was a resource dimension often
partitioned (Luiselli, 2008).
Habitat loss and fragmentation have caused drastic declines in freshwater turtles
(Mitchell and Klemens, 2000). Because habitat quality is vital for population persistence and
important in structuring turtle communities, understanding species-habitat relationships will aid
in assessing fitness and long-term population persistence, criteria essential for conservation
practices (Morrison et al., 2006). To evaluate species-habitat relationships in a sympatric
freshwater turtle community, I assessed habitat partitioning using radio-telemetry data collected
at an isolated preserve within a highly disturbed landscape in northeastern Illinois. The goal of
this project was to determine macro- and micro-habitat use and estimate habitat partitioning and
overlap at both habitat levels among three common and two rare species. My objectives were to
1) evaluate macro- and micro-habitat use for each species 2) compare macro- and micro-habitat
51
use among species 3) measure niche breadth and niche overlap for species at both habitat use
levels and 4) identify partitioning strength of micro-habitat variables among species.
METHODS
Study Site and Species.The study was conducted from May September 2006 at a 124 ha
preserve located in Will County, Illinois and is situated in a matrix of urbanization and industrial
development. The preserve is a prairie-wetland mosaic consisting of various wetland macrohabitats that can be broadly classified as cattail (Typha) marsh, sedge meadow, and pond. The
preserve also lies adjacent to the Des Plaines River and associated riparian macro-habitats such
as scoured backwater ponds and floodplain forest. Micro-habitat characteristics such as
vegetation structure and composition, water depth, canopy cover, and substrate vary substantially
among habitat types and aid in defining the broader habitat categories. For example, presence
and height of emergent vegetation is considerably greater in marsh habitats than in pond or
riparian habitats. Within the preserve boundary, much of the wetland substrate is characterized as
organic; however, substrate within the Des Plaines River and backwater areas is predominantly
characterized as silt. Many transitional areas between habitat types also exist resulting in microhabitat variation within macro-habitat types. During high water events, the river and backwater
pools carry silt into adjacent wetlands within the preserve, altering the substrate composition.
Additionally, within interior wetlands, cattail marsh bordering a sedge meadow typically has
shallower water depths than cattail marsh bordering a pond.
An abundant turtle fauna inhabits the wetland areas within the preserve and the adjacent
riparian habitats (Anthonysamy et al. unpubl.). Common turtle species include the painted turtle
(Chrysemys picta), snapping turtle (Chelydra serpentina), and eastern musk turtle (Sternotherus
52
odoratus); however, two rare turtle species, the Blandings turtle (Emydoidea blandingii) and
spotted turtle (Clemmys guttata) also occur at the preserve. Chelydra serpentina, S. odoratus,
and C. picta are widely distributed and abundant throughout much the United States whereas E.
blandingii and C. guttata have more restricted distributions, are found at lower population
densities, and are considered to be species of conservation concern throughout their range,
mainly because of habitat loss (Ernst and Lovich, 2009).
Field methods.I radio-tracked 61 adult turtles: five male and 15 female E. blandingii, five
male and seven female C. guttata, four male and five female S. odoratus, five male and four
female C. picta, and six male and five female C. serpentina. I affixed radio-transmitters to the
rear marginals of turtles using transmitters and methods as described in Anthonysamy et al. (in
review) and radio-located turtles from three to seven times a week. At each radio-location I
attempted to visually or tactilely confirm presence of the turtle and recorded GPS coordinates
(UTM-NAD 83 CONUS) and a suite of habitat variables.
Macro-habitat use.I plotted turtle location coordinates onto a vegetation community map
provided by the Forest Preserve District of Will County that was field-checked during the study.
Coordinates were assigned to seven macro-habitat categories: cattail marsh, pond, sedge
meadow, river, floodplain (forested and open riparian areas), mesic dolomite prairie, and dry
dolomite prairie. Using the habitat assignments, I calculated the proportion of locations for each
turtle in each habitat and the proportion of available habitat types in the study area. I then used
compositional analysis to assess macro-habitat use vs. availability for each species (Aebischer et
al., 1993). For each turtle, I used the proportion of available and the proportion of used macro-
53
habitats to calculate the difference in log ratios for each macro-habitat pair. To qualitatively
assess macro-habitat use within and among species, differences in log ratios of use vs.
availability between macro-habitat pairs were used to establish rankings in macro-habitat use for
each individual turtle (Aebischer et al., 1993). Rankings ranged from zero to seven (number of
habitat types) with larger ranks representing higher use than smaller ranks. Mean habitat
rankings ( 1 SE) were calculated for each species for each habitat type. To quantitatively assess
differential habitat use among species, I used a multivariate analysis of variance (MANOVA) to
test for differences in log ratio values of use vs. availability among species. Because species
sample size was unequal, I used Gabriels multiple comparison post hoc tests to compare
differences in macro-habitat use between species.
Using macro-habitat proportions, I estimated niche breadth for each species as well as
niche overlap between species. To account for variation in macro-habitat availability, I used the
Proportional Similarity Index (Feinsinger et al. 1981),:
1
0.5
where
= Proportional Similarity Index
= Proportion of radio-locations in macro-habitat i
= Proportion of available macro-habitat i
For broad niche breadths or those where habitats are used in proportion to availability,
Conversely,
= min
= 1.0.
when habitat used is specialized.
Niche overlap in macro-habitat use was calculated between each species pair using the
percentage overlap measure proposed by Renkonen (1938) and given in Krebs (1989) by:

100
54
where

= Percentage macro-habitat use overlap between species j and species k

= Proportion of macro-habitat used i of the total macro-habitat proportions used by
species j and species k

n = Total number of macro-habitats
The percentage overlap measure is interpreted as the area of overlap of resource use between two
species (Krebs, 1989).
Micro-habitat use. I quantified the following micro-habitat structural variables at each radiolocation: structure and type of vegetation, water depth, amount of open water, and substrate type.
I measured water depth at the location of the turtle and height of the tallest plant within 0.5 m of
the turtle. I determined proportion of open water vs. vegetation at the surface by holding a
spherical densiometer upside down above head height (~1.5-2.0 m) and counting the number of
grid dots obscured by water or vegetation to the nearest 1%. I also measured understory canopy
cover (i.e. emergent vegetation, grasses) and overstory canopy cover (i.e. trees) by holding the
densiometer at waist (~ 1.0 m ) and at chest height (~ 1.3 m), respectively. Densiometer
measurements were taken within 0.5 m of the turtle in each cardinal direction and then averaged
across directions. I classified substrate at turtle locations as organic (i.e., unconsolidated with
non-woody debris and a dark color), inorganic (i.e. containing silt, sand, or rock, usually
consolidated and light in color), or mixed and calculated the proportion of locations having
entirely organic substrates for each turtle. Based on published accounts of turtle habitat
associations, I considered organic substrates to indicate higher quality wetlands for the turtle
species in my study (Ross and Anderson, 1990; Kiviat, 1997; Marchand and Litvaitis, 2004).
55
To avoid correlation among micro-habitat variables, I conducted a principle components

analysis (PCA) using the continuous variables from the radio-locations to create new
orthogonally independent variables. Because substrate was categorical variable, it was not
included in the PCA. I chose to include only individuals having at least 20 locations with
complete habitat data in the analyses to ensure adequate sampling and retained components with
eigenvalues > 0.9. For each turtle, I plotted mean component scores against each other to
examine relative micro-habitat niche breadth and niche overlap among species. To identify
patterns of micro-habitat partitioning among species, mean PCA component values and
proportion of locations with organic substrates for each turtle were used in a one-way analysis of
variance (ANOVA) to test for differences in micro-habitat use among species. Proportions were
arcsine-square root transformed prior to analysis. I used Gabriels multiple comparison post hoc
tests to compare differences in micro-habitat use between species. Analyses were conducted in
SPSS 17.0 (SPSS Inc. Chicago, Illinois). Averages are reported as mean 1 S.E and all
significance levels were set at = 0.05.
I used the classification-tree analysis package tree, implemented in R software 2.13.2
(R Development Core Team 2011) to determine how effectively the micro-habitat variables
partitioned the species. Classification trees are non-parametric methods useful for revealing
complex ecological patterns (Death and Fabricius, 2000). The tree was constructed from
principal component scores and proportion of locations with organic substrates of individual
turtles with species as the response variable. Optimal tree size range was identified by using the
cross-validation (cv.tree) code to plot the change in deviance against tree size. I simplified the
tree using the pruning (prune.tree) code to find the tree size closest to five (number of species)
with the lowest misclassification rate. After optimal tree size was determined, I calculated a K
56
statistic to assess tree performance. I calculated K using the method employed by Dellinger et al.
(2007) as follows:

where
#
#
= Ratio of the improvement of the optimal tree classification from chance

classification and a tree with perfect classification
= # actual observations correctly classified by tree

= # observations correctly classified by chance on average
= # observations correctly classified by a perfect tree
I used the benchmark ranges for values of K created by Landis and Koch (1977) to evaluate
strength of the optimal tree: < 0.00 poor, 0.00-0.20 slight, 0.21-0.40 fair, 0.41-0.60 moderate,
0.61-0.80 substantial, and 0.81-1.00 almost perfect.
RESULTS
Fifty turtles had at least 20 radio-locations with complete habitat data and were included in the
analyses: five male and 13 female E. blandingii, five male and five female C. guttata, four male
and five female S. odoratus, four male and two female C. picta, and four male and three female
C. serpentina (Appendix F). Average number of radio-locations for individuals used in analyses
was 75.5 5.55 for E. blandingii, 69.0 2.56 for C. guttata, 41.8 4.83 for C. picta, 66.4 4.25
for C. serpentina, and 50.7 5.52 for S. odoratus. Of the 50 turtles retained for analysis, one
male E. blandingii, one male C. guttata and one female C. picta, were depredated during the
study. Use of dry dolomite prairie was minimal for all species and will not be considered
further.
57
Macro-habitat use.Qualitative assessments of wetland macro-habitat use vs. availability

differed substantially resulting in variation in mean macro-habitat ranks among species (Table
2.1, Fig. 2.1). In relation to availability, E. blandingii and C. guttata most often used marshes,
whereas C. picta, C. serpentina, and S. odoratus most often use ponds. Among wetland macrohabitats, floodplain was used the least among all species. The most notable differences in mean
rankings were between C. guttata and the common species; C. guttata used mesic dolomite
prairie, marsh, and sedge meadow to a greater extent whereas C. picta, C. serpentina, and S.
odoratus used river and pond to a greater extent (Fig. 2.1). For mesic dolomite prairie, river,
marsh, sedge meadow, and pond macro-habitats, E. blandingii ranked intermediately between C.
guttata and the common species.
The results of the MANOVA also showed that proportional use of macro-habitats
differed among species (Wilks = 0.163, F24, 140 = 3.996, P < 0.001). Post-hoc tests were
consistent with qualitative measures of the macro-habitat rankings (Appendix G). Mesic prairie
was used more by C. guttata than C. picta, C. serpentina, and S. odoratus (P < 0.012). Further,
C. guttata also used sedge meadow more than S. odoratus (P = 0.007). Both C. serpentina and S.
odoratus used river more than C. guttata (P < 0.045). Finally, C. picta, C. serpentina, and S.
odoratus used pond to a greater extent than E. blandingii and C. guttata (P < 0.008). No
significant differences in macro-habitat use were detected between the two rare species or among
the three common species.
Macro-habitat niche breadth was broadest for E. blandingii (0.56) followed by C.
serpentina (0.52), C. guttata (0.34), C. picta (0.32), and S. odoratus (0.20). Niche overlap of
macro-habitat use was greatest among the common species and lowest between C. guttata and
58
the common species (Table 2.2). Emydoidea blandingii shared intermediate levels of overlap
with C. guttata and C. serpentina and lower levels of overlap with C. picta and S. odoratus.
Micro-habitat use.Two components were retained from the PCA analysis of micro-habitat
variables recorded at turtle radio-locations. The first component (PC1) explained 58% of the
variance. The variables loading high on PC1 were vegetation surface cover, vegetation height,
and understory canopy cover (positive) and water depth and water surface cover (negative; Table
2.3). The second component (PC2) explained 18% of the variance. Overstory canopy cover
(negative) was the only variable to load high on PC2 (Table 2.3). Plots of mean component
scores illustrated narrower dimensions of micro-habitat use for C. guttata, C. picta, and S.
odoratus compared to E. blandingii and C. serpentina (Fig. 2.2). Further, separation of C. guttata
from C. picta and S. odoratus along gradients of vegetation and water characteristics (PC1 axis)
was apparent, whereas micro-habitat use of E. blandingii and C. serpentina overlapped with
multiple other species (Fig. 2.2).
For the ANOVA, PC1, water and vegetation characteristics (F4, 45 = 29.40, P < 0.001),
PC2, overstory canopy cover (F4, 45 = 3.93, P = 0.008), and substrate (F4, 45 = 17.14, P < 0.001)
differed significantly among species. Post hoc tests revealed that micro-habitat use of C. guttata
was characterized by shallower water depths, taller vegetation heights, higher vegetation surface
cover, greater amount of understory cover, and more organic substrates than all other species (P
0.016; Fig. 2.3; see Appendix G). Similarly, micro-habitat use of E. blandingii was
characterized by shallower water depths, and greater vegetation structure and organic substrates
than C. picta and S. odoratus (P < 0.001) but not C. serpentina. No differences in water and
vegetation or substrate micro-habitat characteristics were detected among the common species.
59
Micro-habitat use of shoreline tree cover was greater for S. odoratus than E. blandingii and C.
picta (P 0.032).
The classification tree analysis most strongly differentiated species by PC1 (water and
vegetation characteristics) followed by PC2 (shoreline tree cover; Fig. 2.4). Optimal tree size
derived from cross-validation and pruning consisted of four terminal nodes, one for each species
except C. serpentina. Higher PC1 values ( 0.39), or use of more highly vegetated micro-habitats
with less water (i.e. shallow cattail marsh), most strongly differentiated C. guttata from all other
species. Further, moderate use of micro-habitats with more vegetation and less water ( -0.58)
differentiated E. blandingii from S. odoratus and C. picta. Lastly, use of micro-habitats with
greater shoreline tree cover separated S. odoratus from C. picta. Substrate was not selected by
the tree package for tree construction presumably because substrate use was correlated with
PC1 and rendered no additional information. The optimal tree had an overall correct
classification rate of 0.70 and correctly classified 100% of C. guttata, 83% of E. blandingii, 0%
of C. serpentina, 83% of S. odoratus, and 83% of C. picta. Three E. blandingii were
misclassified as C. guttata. One S. odoratus was misclassified as C. picta and vice versa. One C.
serpentina was misclassified as an S. odoratus and the remaining six individuals were
misclassified as E. blandingii. The classification tree demonstrated substantial agreement (K
statistic = 0.62) of the tree model based on the benchmark range of Landis and Koch (1977).
DISCUSSION
Macro-habitat analysis was useful for identifying coarse patterns of habitat use and partitioning
in my study. Emydoidea blandingii, C. guttata, C. serpentina, S. odoratus, and C. picta are
known to inhabit a variety of wetland habitats throughout their ranges but E. blandingii and C.
60
guttata are less tolerant of habitat degradation (Ernst and Lovich, 2009). In this study, all species
used multiple macro-habitat types, but the rare turtle species, E. blandingii and C. guttata, most
frequently used cattail marsh macro-habitats whereas the common species (C. picta, C.
serpentina, and S. odoratus) most frequently used pond macro-habitats. Emydoidea blandingii
used the highest number of macro-habitat types (N = 7) followed by C. serpentina (N = 5) and C.
guttata (N = 4) whereas S. odoratus and C. picta used the fewest number of macro-habitats (N =
3). Use of multiple habitat types at a study site has also been documented for other populations
of E. blandingii and C. guttata (Joyal et al., 2001; Edge et al., 2010). In my study, cattail marsh
was the most available wetland habitat and was the only macro-habitat used by all species.
The quantitative comparison of macro-habitat use between species revealed that use of
mesic prairie, sedge meadow, river, and pond macro-habitats differed between C. guttata and
common species while only use of pond macro-habitats differed between E. blandingii and two
common species, S. odoratus, and C. picta. Similarly, Bury and Germano (2003) found that
within turtle communities in Nebraska, E. blandingii occurred most often in marshes and small
ponds whereas more C. picta occurred in lakes and open waters. Although I failed to detect
differences in macro-habitat use between E. blandingii and C. guttata, differences in seasonal
patterns of macro-habitat use between these species have been observed in Maine (Joyal et al.,
2001). Joyal et al. (2001) reported that use of permanent pools was greater in E. blandingii and
C. guttata used wet meadows whereas E. blandingii did not.
I found that species most strongly partitioned micro-habitat along an axis comprised of
water depth, water and vegetative surface cover, vegetation height, and understory canopy cover.
Clemmys guttata and S. odoratus displayed a narrower range of use of vegetative and water
characteristics compared to the other species; however, differentiation in water and vegetation
61
micro-habitat use was greatest between C. guttata and all common species. Separation of C.
guttata in micro-habitat use from the other species was also supported by the classification tree
analysis. Similarly, water depth and vegetation characteristics were also partitioned in different
size classes of juvenile E. blandingii in Minnesota (Pappas and Brecke, 1992) and vegetation
structure and open water affected habitat selection of adult E. blandingii in Ontario (Millar and
Blouin-Demers, 2011). Water characteristics such as depth, open water, and velocity have been
key determinants of habitat use in other freshwater turtles (Plummer, 1977; Souza and Abe,
1998). Proportion of organic substrates at radio-locations also differentiated habitat use among
species in this study; use of organic substrates was highest among C. guttata and E. blandingii.
Similarly, substrate characteristics were shown to be important for differentiating habitat use
among species of map turtles, Graptemys sp. (Fuselier and Edds, 1994).
Micro-habitat use differentiated species to a greater extent than macro-habitat use
indicating that species were using distinct micro-habitats within macro-habitats. For example, no
difference in macro-habitat use was detected between the two rare species (both highly used
cattail marsh) yet C. guttata used shallower wetlands with more vegetation structure and organic
substrates than E. blandingii. The interior wetlands at my study site contained more organic
substrates and likely provided higher quality habitat compared to the peripheral preserve areas
that are subjected to flooding and silt deposition by the Des Plaines River. I observed that C.
guttata and E. blandingii most often used these higher quality interior cattail marsh habitats;
however, some E. blandingii occasionally used peripheral wetlands and shallow areas of the Des
Plaines River. For example, three E. blandingii (EMBL 7, EMBL 22, & EMBL 36) used the
river > 50% of the time whereas C. guttata almost never used silted peripheral wetlands and
were never observed in the river. Further, E. blandingii are obligated to seek refuge and use the
62
river and surrounding riparian habitats more extensively during years of drought when interior
marsh habitat becomes dry (Anthonysamy et al. in review). Similarly, Fuselier and Edds (1994)
found that finer scale environmental variables differentiated three Graptemys species even
though overlap in habitat use was high.
Although C. picta and S. odoratus exhibited high overlap in macro-habitat use and
similar micro-habitat use of vegetation and water characteristics, S. odoratus were more apt to
use micro-habitats near the shore as they occasionally used mammal excavations and undercuts
within the bank. Use of muskrat burrows by S. odoratus has also been documented by Ernst
(1986). Thus, greater use of shoreline tree cover by S. odoratus than C. picta is not necessarily a
preference for shaded habitats but more likely a preference for a different resource characteristic
(e.g. foraging, basking, dietary) that is coincidently associated with floodplain habitat such as
riparian forest that often bordered macro-habitats used by these species.
Measures of niche breadth and niche overlap also varied among species. Among all
species, E. blandingii and C. serpentina most broadly and similarly used macro- and microhabitats and maintained a relatively large measure of niche breadth. Further, these two species
also demonstrated a considerable amount of niche overlap with the other species in their
respective rare and common species groups; E. blandingii with C. guttata and C. serpentina with
C. picta and S. odoratus. Hence, these findings indicated that E. blandingii and C. serpentina
were functioning as habitat generalists. Swihart et al. (2006) also found that C. serpentina in
Indiana had the greatest niche breadth among a group of eight turtle species including E.
blandingii, S. odoratus, and C. picta. Interestingly, in my study, C. picta and S. odoratus
exhibited the narrowest measures of macro-habitat niche breadth, but attained the highest
measure of niche overlap (82.9; Table 2.2) and used the most silted and peripheral habitats.
63
Further, micro-habitat use of water and vegetation structure of these species overlapped
substantially with C. serpentina. Finally, C. guttata demonstrated a narrower but intermediate
range of macro-habitat niche breadth compared to the other species; however use of microhabitat was most divergent for this species and was also restricted to the higher quality, interior
wetlands, with organic substrates. These findings suggest that C. guttata is a micro-habitat
specialist. Nevertheless, my estimates of macro-habitat use and niche breadth measures should
be interpreted with caution as these measurements were calculated based on the proportion of
available macro-habitats as delineated by me and sample size was limited for some species.
Resource partitioning may result from competition, predation, and physiological
constraints, as well as complex interactions among these biological mechanisms (Toft, 1985). In
a review of resource partitioning among freshwater turtles, Luiselli (2008) concluded that
partitioning was most likely a result of interspecific competition. Competition and aggressive
behavior have been documented among emydid turtle species for basking sites (Lovich, 1988;
Lindeman, 1999; Cadi and Joly, 2003). In addition, differential survival (Cadi and Joly, 2004)
and differential growth in low resource conditions (Aresco, 2010) have been observed between
species. In my study, interspecific competition for resources should be greatest among the
species with the greatest niche overlap, C. serpentina, S. odoratus, and C. picta; however, I did
not observe competitive interactions or aggressive behaviors among turtle species.
Predation is a critical threat to turtles at my study site as one male E. blandingii, one male
C. guttata and one female C. picta, were depredated during this study. Turtles exhibit patterns of
size-dependent predation with smaller body sizes being more susceptible to predators (Janzen,
1993; Congdon et al., 1993; 1994; Tucker et al., 1999; Janzen et al., 2000). My findings
supported this idea in that the second smallest species, C. guttata, was the least aquatic and
64
strictly used micro-habitats with higher amounts of vegetation structure that afforded more
protection from predation than more open water habitats. However, the smallest species in this
study, and possibly the most aquatic, S. odoratus, used deeper wetlands with little to no
vegetation cover and overlapped in habitat use with the largest species, C. serpentina. Hence, the
predation risk/body size association may be influencing habitat use in turtle species at my study
site but other factors are probably also contributing to differential habitat use among species.
Habitat partitioning observed in this study is likely related to species-specific traits. The
species in this study exhibit variation in traits such as morphometrics, foraging strategies, dietary
preferences, and basking habits (see Ernst and Lovich, 2009) that have been shown to influence
habitat partitioning in turtles (Plummer, 1977; Vogt, 1981; Williams and Christiansen, 1981;
Hart, 1983; Vogt and Guzman, 1988; Lindeman, 2000). Because different habitat types likely
vary in food availability, thermal properties, and ease of maneuverability, use of habitat types
that optimize fitness should also be expected to differ among species. Compared to C. guttata, S.
odoratus and C. picta have evolved morphological characteristics such as extensive toe-webbing
that improve aquatic locomotion in deeper, more open water habitats with less vegetation
(Ludwig et al., 2007) that helps to explain the strong divergence in habitat use observed between
these species. In another scenario, dense cattail stands and shallow wetlands may inhibit foraging
in larger species with higher energetic demands (i.e. C. serpentina) but may provide optimal
refugia and foraging opportunities for small species (i.e. C. guttata). Emydoidea blandingii and
C. serpentina tended to use a greater range of habitats than smaller species, and presumably
because of their larger body size, likely exploited larger-sized dietary items compared to the
smaller species (Costa et al., 2008). In addition, frequency and method (e.g. aerial, surface, land)
of basking varies for the species in this study (Ernst and Lovich, 2009; pers. obs.); therefore
65
species may have also used micro-habitat features that were conducive for species-specific
basking habits.
Conservation Implications.Species-habitat relationships and dimensions of habitat

partitioning in sympatric turtle communities are important components for the conservation and
management of freshwater turtles. I emphasize the need to assess fine-scale micro-habitat use
because species with high overlap in macro-habitat use showed distinct differences when microhabitat variables were included. Additionally, management efforts for sympatric species of
conservation concern should be considered for each species independently. For example, the two
rare species demonstrated different overall patterns of habitat use; C. guttata was more of a
habitat specialist whereas E. blandingii was more of a habitat generalist. Species that are habitat
specialists are predicted be less tolerant of wetland loss and degradation than those that are
habitat generalists (Henle et al., 2004; Ewers and Didham, 2006). Compared to the other turtle
species C. guttata is most vulnerable to degradation of high quality interior shallow cattail
marsh, sedge meadow, and mesic dolomite prairie from siltation caused by flooding of the Des
Plaines River. Illinois populations of C. guttata represent the western-most periphery of this
species distribution which further increases these populations vulnerability to habitat
fragmentation (Swihart et al., 2006). For E. blandingii, these findings are surprising as this
species is threatened throughout its range due to habitat loss (Ernst and Lovich, 2009); however,
E. blandingii is highly vagile and capable of long-distance movements (Rowe and Moll, 1991;
Sexton, 1995; Piepgras and Lang, 2000; Joyal et al., 2001, Chapter One) that allow it to access a
greater number of wetlands such as the river and peripheral pond habitats in my study. These
66
findings suggest that multiple macro-habitat types and wide variation in water and vegetation
micro-habitat characteristics are necessary to support a diverse freshwater turtle community.
67
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73
TABLES
74
Table 2.1 Proportions of available macro-habitat used by ten C. guttata (CLGU), 18 E.

blandingii (EMBL), seven C. serpentina (CHSE), nine S. odoratus (STOD), and six C. picta
(CHPI) radio-located at a preserve in Will County, Illinois during 2006.
Proportion Available
Macro-habitat
Mesic Prairie
Dry Prairie
Floodplain
River
Marsh
Sedge Meadow
Pond
Proportion Used
CLGU EMBL
0.11
0.25
0.20
0.16
0.23
0.04
0.02
0.06
0.01
0.00
0.00
0.81
0.12
0.00
0.01
0.01
0.09
0.21
0.51
0.06
0.10
CHSE
STOD
CHPI
0.00
0.00
0.10
0.15
0.23
0.03
0.50
0.00
0.00
0.00
0.12
0.06
0.00
0.82
0.00
0.00
0.00
0.07
0.22
0.00
0.70
75
Table 2.2 Macro-habitat niche overlap values for ten C. guttata (CLGU), 18 E. blandingii
(EMBL), seven C. serpentina (CHSE), nine S. odoratus (STOD), and six C. picta (CHPI) radiolocated at a preserve in Will County, Illinois during 2006. Measures of niche breadth for each
species are underlined and appear on the diagonal.
Species
CLGU
EMBL
CHSE
STOD
CHPI
CLGU EMBL
0.34
59.7
25.5
5.7
22.8
0.56
59.5
28.0
39.3
CHSE
STOD
CHPI
0.52
67.9
79.3
0.20
82.9
0.32
76
Variable
% Water Surface Cover
% Vegetation Surface Cover
Vegetation Height (cm)
Water Depth (cm)
% Understory Canopy Cover
% Overstory Canopy Cover
PC1
-0.944
0.933
0.769
-0.745
0.619
0.062
PC2
-0.083
0.127
0.437
0.229
0.571
-0.831
Loadings
Coefficients
PC1
PC2
-0.293
0.330
-0.305
0.094
0.294
-0.053
0.167
-0.734
0.110
0.388
0.184
0.246
77
Table 2.3 The coefficients and loadings on principal component one (PC1) and two (PC2) retained for micro-habitat use variables
collected for ten C. guttata, 18 E. blandingii, seven C. serpentina, nine S. odoratus, and six C. picta radio-located at a preserve in Will
County, Illinois during 2006.
FIGURES
78
Fig. 2.1 Mean wetland macro-habitat rankings derived using compositional analysis for ten C.
guttata (CLGU), 18 E. blandingii (EMBL), seven C. serpentina (CHSE), nine S. odoratus
(STOD), and six C. picta (CHPI) radio-located at a preserve in Will County, Illinois during 2006.
8.0
CLGU
EMBL
CHSE
STOD
CHPI
Mean Habitat Rank
7.0
6.0
5.0
4.0
3.0
2.0
1.0
0.0
Mesic Floodplain
Dolomite
Prairie
River
Cattail
Marsh
Sedge
Meadow
Pond
Habitat Type
79
Fig. 2.2 Plot of mean pricipal component scores (PC1 vs PC2) calculated from micro-habitat
variables collected at radio-locations for ten C. guttata (CLGU), 18 E. blandingii (EMBL), seven
C. serpentina (CHSE), nine S. odoratus (STOD), and six C. picta (CHPI) radio-located at a
preserve in Will County, Illinois during 2006. Polygons connect outermost points and illustrate
relative micro-habitat niche breadth size and niche breadth overlap among species.
CLGU
EMBL
CHSE
STOD
CHPI
Less Overstory
Canopy Cover
1.5
PC 2
1.0
0.5
PC 1
0.0
-2.0
-1.5
Deep Water
Less Vegetation
-1.0
-0.5
0.0
-0.5
0.5
1.0
1.5
2.0
Shallow Water
More Vegetation
-1.0
More Overstory
Canopy Cover
-1.5
80
D)
0.0
10.0
20.0
30.0
40.0
50.0
0.00
0.10
0.20
0.30
0.40
0.50
0.60
0.70
0.80
CLGU EMBL CHSE STOD CHPI
0.45
0.40
0.35
0.30
0.25
0.20
0.15
0.10
0.05
0.00
E)
1.00
0.90
0.80
0.70
0.60
0.50
0.40
0.30
0.20
0.10
0.00
B)
Proportion of Vegetation
Cover
Proportion of
Understory Cover
A)
Species
C)
F)
0.00
0.05
0.10
0.15
0.20
0.25
180.0
160.0
140.0
120.0
100.0
80.0
60.0
40.0
20.0
0.0
81
Fig. 2.3 Mean micro-habitat values and standard errors of A) proportion of water surface cover, B) proportion of vegetation surface
cover, C) vegetation height, D) water depth, E) proportion of understory canopy cover, G) proportion of overstory canopy cover and
F) proportion of locations having organic substrates for ten C. guttata (CLGU), 18 E. blandingii (EMBL), seven C. serpentina
(CHSE), nine S. odoratus (STOD), and six C. picta (CHPI) radio-located at a preserve in Will County, Illinois during 2006.
Proportion of Water
Cover
Water Depth (cm)
Vegetation Height (cm)

Proportion of
Overstory Cover
F)
1.00
0.90
0.80
0.70
0.60
0.50
0.40
0.30
0.20
0.10
0.00
Species
Fig. 2.3 (cont.)
Frequency of Organic
Substrate
82
Fig. 2.4 Classification tree based on micro-habitat measures collected from radio locations for C.
guttata (CLGU), E. blandingii (EMBL), C. serpentina (CHSE), S. odoratus (STOD), and C.
picta (CHPI) radio-located at a preserve in Will County, Illinois during 2006. Increasing positive
values for PC1 represent microhabitats with less water and more vegetation. Increasing positive
values for PC2 represent microhabitats with less shoreline overstory canopy cover. The length of
the vertical line below each split indicates variable importance in the separation. Sample size and
species composition of resulting classification for each node is shown.
Species
CLGU
EMBL
CHSE
STOD
CHPI
Total
N
10
18
7
9
6
50
PC1
< 0.39
0.39
PC1
< -0.58
-0.58
CLGU
N = 13
CLGU 77%
EMBL 23%
PC2
< -0.02
-0.02
EMBL
N = 25
STOD
N=6
CHPI
N=6
CHSE 17%
STOD 83%
STOD 17%
CHPI 83%
EMBL 60%
CHSE 24%
STOD 12%
CHPI 4%
83
CHAPTER 3
COMPARISON OF POPULATION GENETIC STRUCTURE AMONG THREE
SYMPATRIC FRESHWATER TURTLE SPECIES
INTRODUCTION
Anthropogenic landscape fragmentation results in small, isolated, remnant populations
vulnerable to decreased levels of genetic diversity via genetic drift and reduced gene flow
(Spradling et al. 2010, Reed et al. 2011). In many cases, this is compounded by increased levels
of inbreeding. Loss of genetic diversity and inbreeding can lead to reduced fitness from the
expression of deleterious genes and compromise survival, fertility, and general health
(Westemeier et al. 1998) as well as impair the ability of populations to adapt to a changing
environment (Willi et al. 2006).
Comparing genetic structure in sympatric species of similar taxa that vary in life history
and ecological traits improves our understanding of how species respond to fragmentation
(Steele et al. 2009, DiLeo et al. 2010, Goldberg and Waits 2010). Variation in species-specific
traits such as dispersal ability, reproductive effort, and ecological specialization influences
genetic processes among species. For example, three sympatric snake species that varied in body
size and vagility exhibited marked differences in gene flow and genetic population structure in a
subdivided island/mainland system (King and Lawson 2001). In turtles, lack of dispersal can
result in the loss of gene flow between populations (Kou and Janzen 2004, Richtsmeier et al.
2008), and might ultimately lead to reduced genetic variation (Gray 1995, Parker and Whiteman
1993).
In this study, I examined genetic diversity and genetic divergence in three sympatric
freshwater turtle species sampled from three fragmented and one intact site in Illinois. The
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species represent two families; Emydidae [Blandings turtle (Emydoidea blandingii), painted
turtle (Chrysemys picta)] and Chelydridae [common snapping turtle (Chelydra serpentina)] and
are of different conservation status (Ernst & Lovich 2009). These three species vary in a number
of characteristics such as life history traits (Congdon et al. 1993, Congdon et al. 1994, Congdon
et al. 2003, McGuire 2011, McGuire et al. 2011), vagility (Chapter One), and habitat use
(Chapter Two), which can influence gene flow and loss of genetic diversity in a fragmented
landscape. For example, compared to C. picta and C. serpentina, E. blandingii has lower
reproductive output (clutch size, annual clutch frequency) as well as a longer generation time
(Congdon et al. 1993, Congdon et al. 1994, Congdon et al. 2003, McGuire 2011, McGuire et al.
2011). In addition, C. picta and C. serpentina are widely distributed and abundant throughout
much of the United States, whereas E. blandingii has a more restricted distribution and is
considered rare throughout much of its range (Ernst & Lovich 2009).
I employed microsatellite DNA markers in three turtle species across four study sites to
investigate the effects of fragmentation and species-specific differences in ecological/ life history
traits on and genetic diversity and genetic divergence. I tested the following predictions: 1) All
species will have decreased levels of genetic diversity in the fragmented sites compared to the
intact site; 2) In the fragmented sites, E. blandingii will have lower levels of genetic diversity
and higher levels of genetic divergence within and among species compared to the common
species (C. picta and C. serpentina); 3) All species from the fragmented sites will show evidence
of recent population bottlenecks; 4) Future levels of genetic diversity would be lower for E.
blandingii than C. picta or C. serpentina. Predictions 2 and 4 stem from the lower reproductive
output, longer generation time, and lower population size of E. blandingii (Ernst and Lovich,
2009). Finally, because females of many species of turtles are philopatric to nesting locations
85
(Congdon et al. 1983, Congdon et al. 1987, Valenzuela & Janzen 2001, Rowe et al. 2005), I
predict lower levels of gene flow in females compared to males, for all species.
METHODS
Study sites
My study was conducted within the Lower Des Plaines River Valley (LDPRV) in
northeastern Illinois. This area was once a prairie-dominated landscape (Bowles & McBride
2001) composed of semi-contiguous prairie-wetland matrices that allowed turtles to freely
disperse along the river corridor without anthropogenic impediment. However, since the early
1800s there have been drastic environmental changes as a result of European settlement and
associated anthropogenic alterations. Gradually, over the past 150 years, agriculture, shipping
canals, railways, roadways, quarries, industrial parks, and towns have come to dominate the
landscape. Remaining natural areas are effectively isolated from one another except for their
connection along the narrow Des Plaines River riparian zone.
Turtles were sampled at four sites along the LDPRV; three small, isolated sites in Will
County (Will 1-3); and one large, more intact site in Grundy County (Grundy; Fig. 3.1). The sites
are located along a 40 km stretch of the Des Plaines River. Will 1 (95 ha) and Will 2 (188 ha) are
separated by 1 km, and Will 2 and Will 3 (124 ha) by 6 km. The Grundy site (1247 ha) is the
largest remnant prairie in Illinois, and is located near the confluence of the Des Plaines River and
Kankakee rivers, approximately 34 km by river southwest of Will 3. The LDPRV sites are
composed of a prairie-wetland matrix that is inhabited by a diverse turtle assemblage with the
southernmost site (Grundy) providing habitat for large and presumably genetically diverse turtle
populations (Banning et al. 2006, Dreslik et al. 2010, Dreslik et al. 2011).
86
DNA extraction
I collected tissue samples from adult E. blandingii, C. picta, and C. serpentina, captured
during trapping and radio-telemetry surveys conducted from 2004 2009. Blood (0.1-0.3 cc)
was collected from the sub-carapacial sinus (Fisher 2003) of live turtles using a 25 gauge
needle and 1 cc syringe. Tail clips and liver tissue were taken from dead turtles found on roads at
the study sites. I preserved tissues in 95% ethanol or Queens lysis buffer (Seutin et al. 1991) and
stored samples at -80C until DNA extraction. I extracted whole genomic DNA from tissue
samples using the Qiagen DNeasy Blood & Tissue Kit (QIAGEN INC.) following the
manufacturers protocol, with the exception that I digested tissue samples overnight in the
proteinase K solution.
DNA amplification, linkage disequilibrium, and Hardy-Weinberg equilibrium

For E. blandingii and C. picta, I screened 21 microsatellite loci using primers developed
for E. blandingii ([BTCA9; Libants et al. 2004] [Eb09, Eb17, Eb19; Osentoski et al. 2002]) and
bog turtle (Glyptemys muhlenbergii; GmuD21, GmuD55, GmuD70, GmuD87, GmuD90,
GmuD93, GmuD121, GmuB08, GmuA18, GmuA19, GmuA32; King and Julian 2004). For C.
serpentina, I screened nine microsatellite loci using primers developed for alligator snapping
turtle (Macrochelys temminckii; MteA105, MteB103, MteC1, MteC112, MteD2, MteD9,
MteD106, MteD109, MteD111; Hackler et al. 2007). Based on the results of initial primer
testing, I grouped favorable primers into multiplex panels (groups of fluorescent dye-labeled
primers that successfully amplify target DNA regions under similar conditions using polymerase
chain reaction [PCR)]). I determined that 15 primers amplified target DNA in E. blandingii and
C. picta samples. I grouped those primers into four multiplex panels (Appendix H). Seven
87
primers amplified target DNA in C. serpentina samples and were grouped into two multiplex
panels (Appendix H).
I conducted PCR for all panels in 10 l volumes using 0.2-0.9 mM of each primer, 1X
GoTaq Flexi buffer, 2.5-5.0 mM MgCl2, 0.2 mM dNTP, 0.5-1.0 U of Flexi GoTaq DNA
polymerase (Promega), and 1.0 l template DNA. Multiplex reactions were carried out under the
following conditions: initial denaturation at 95 oC for 3 min, followed by 15 cycles of 95 oC for
45 s, a panel-specific annealing temperature for 45 s, and a 72 oC elongation for 30 s, followed
by an additional 25 cycles of 95 oC for 30 s, a panel-specific annealing temperature for 30 s, and
a 72 oC elongation for 15 s, followed by a final extension at 72 oC for 20 min.
Fragment analysis of resulting PCR products was carried out on an automated Applied
Biosystems (ABI) Prism 3730xl sequencer at the W. M. Keck Center at the University of Illinois,
Champaign. An internal size standard (Liz 500) was run with each sample and I scored alleles
using GENEMAPPER 4.1 software (ABI). Within each species, I identified possible null alleles,
large allele dropout, and scoring errors due to stutter peaks using MICRO-CHECKER 2.2.3 (van
Oosterhout et al. 2004). For each species at each study site I tested for linkage disequilibrium
(Markov Chain parameters: 10000 dememorisation steps, 500 batches, 5000 iterations) between
all pairs of loci and tested for departures from Hardy-Weinberg equilibrium (HWE) for each
locus using exact tests in GENEPOP 4.0 (Rousset 2008). Sequential Bonferroni correction was
used to control for multiple comparisons (Rice 1989).
Genetic diversity within species across sites

For each species and site, I estimated allele frequencies, observed heterozygosity (Ho),
expected heterozygosity (He), and inbreeding coefficients (FIS) using GENALEX 6.41 (Peakall &
88
Smouse 2006). In HP-RARE v. June-6-2006 (Kalinowski 2005) I calculated allelic richness (AR)
and private allelic richness (PAR), measures of genetic diversity derived from rarefaction and
corrected for variable sample sizes. I used a paired Wilcoxon rank sum test in SPSS 17.0 (SPSS
Inc. Chicago, Illinois)to test for differences in the amount of genetic diversity (i.e. AR, PAR, Ho)
in each species between the intact Grundy County and each of the fragmented Will County sites.
Genetic divergence within species

To assess genetic divergence among sites, I conducted pairwise FST analysis (999
permutations, interpolated missing data) and an analysis of molecular variance (AMOVA) in
GENALEX. In addition, I used the Bayesian clustering method implemented in program
STRUCTURE 2.3.3 (Pritchard et al. 2000) to further assess genetic structure among sampling
locations. I tested two simulations, one without and one with prior sampling location information
(LOCPRIOR) to assist clustering and assess levels of migration between sites (Pritchard et al.
2000). For remaining parameters, I selected the admixture ancestry model and the correlated
allele frequency model parameter options for both simulations. Five replicate analyses were run
for K values ranging from 1 to 4 (number of sampling locations) using a specified burn-in length
of 500,000 iterations followed by 1,000,000 Markov Chain Monte Carlo (MCMC) replicates. I
assumed no substructure in the intact Grundy County site. I determined the optimal number of
clusters for each simulation by using the online software STRUCTURE HARVESTER 0.6.1 (Earl &
vonHoldt 2011) to calculate ad hoc statistic K described by Evanno et al. (2005).
89
Genetic divergence among species

To compare genetic divergence among the three species, I averaged two standardized
measures of genetic divergence GST (Hedrick 2005, Ryman & Leimar 2009) and Dest (Jost 2008)
for each species across sites. These measures allow for comparisons between species with
different numbers of and variability among loci (Hedrick 2005, Jost 2008) and have been used in
recent studies to compare divergence in sympatric species of salamanders (Steele et al. 2009)
and bumble bees (Lozier et al. 2011). Both GST and Dest were estimated with 95% confidence
intervals (CIs) using 1000 bootstrap repetitions in the R package DEMEtics (Gerlach et al.
2010) implemented in R software 2.13.2. Significance was determined by the non-overlap of
95% CIs.
Sex-biased dispersal
Previous studies have documented nest-site fidelity of adult females in the turtle species
used in this study (Congdon et al. 1983; Congdon et al. 1987; Valenzuela and Janzen 2001;
Rowe et al. 2005). I assessed the presence of sex-biased gene flow among sites using the biased
dispersal option in FSTAT V. 2.9.3.2 (Goudet 1995). I tested for differences in the mean
assignment indices (mAIc), the variance in assignment indices (vAIc), FST, and FIS between
males and females (1000 permutations; Goudet 2002). To compare the potential impacts of
fragmentation on gene flow patterns, I conducted tests for two scenarios; across all sites and
across fragmented sites only. If males are dispersing more than females and sites consist of both
resident and migrant males but mostly resident females, then males should have a negative mAIc
whereas females should have a positive mAIc (Goudet et al. 2002). In addition, males should
exhibit larger vAIc values than females and pairwise FST among sites should be greater for
90
females than males. Finally, measures of FIS should be higher in males because sites should
consist of both resident and migrant males, indicating a heterozygote deficiency (i.e. Wahlund
effect; Goudet et al. 2002).
Bottlenecks
I examined sites for loss of genetic diversity using two different tests in the program
BOTTLENECK 1.2.02 (Piry et al. 1999). For historically recent bottlenecks (0.2 - 4 Ne
generations), I tested whether the observed heterozygosity was higher than expected under the
assumption of mutation-drift equilibrium (Luikart & Cornuet 1998) using the two-phase
mutation model (TPM) option. This model consists of a combination of single step and multiple
step mutations, as recommended for microsatellite data (Di Rienzo et al. 1994, Piry et al. 1999)
and the TPM mutation pattern has been observed in microsatellites documented in sea turtles
(Hoekert et al. 2002). To test for historic population declines, I used the Wilcoxon sign test
(Cornuet & Luikart 1996, Luikart & Cornuet 1998) to test for an excess of heterozygosity at each
study site. The TPM model consisted of 95% single steps and 5% multiple steps with variance
for mutation size set to 12 as recommended by Piry et al. (1999). Further, I tested for the effects
of alterations in these parameters in the model by varying the frequency of single step (98%,
90%) and multiple step mutations (2%, 10%) in two additional scenarios (Rivalan et al. 2006).
To test for more recent population declines (few dozen generations), I used a qualitative mode
shift test (Luikart et al. 1998) to evaluate shifts in allele frequencies from loss of rare alleles. The
input file for C. picta failed to run in program BOTTLENECK when the data set included the locus
GmuD70; thus this locus was excluded from Wilcoxon sign test and mode-shift test for this
species.
91
I also assessed historic bottleneck effects using the M-ratio method of Garza &
Williamson (2001). This method is used to detect bottlenecks by comparing the mean ratio of the
number of alleles to the range in allele size under the TPM model and essentially measures the
gaps between the largest and smallest allele, which would be larger in sites that had
experienced genetic drift. Loss of alleles in a bottlenecked population would produce a smaller
ratio compared to a population under mutation-drift equilibrium (Garza & Williamson 2001). I
tested for significance in M values for each locus across each site by comparing estimated values
of M to critical values of M (Mc) using the software programs M_P_VAL.EXE and
CRITICAL_M.EXE (Garza & Williamson 2001). Both programs require three input parameters to
estimate M values: percentage of single-step mutations (ps), average size of non-stepwise
mutations (g), and a population specific (4Neu) where Ne is effective population size and u is
the mutation rate. I used ps = 0.9 and g = 3.5 as suggested by Garza & Williamson (2001) but
because pre-bottleneck population size was unknown, I tested for values ranging from 0.1 to 10
(e.g. Busch et al. 2007, Parga et al. 2012).
Future loss of genetic diversity

To predict and compare future loss of genetic variation among species from genetic drift,
I used the program BOTTLESIM v.2.6 (Kuo & Janzen 2003) to simulate levels of allelic diversity
and heterozygosity remaining over a 300-year period. This program includes a scenario for longlived species with overlapping generations (i.e. turtles) and requires input of life history trait and
demographic parameters such as longevity, age of maturity, mating system, population size, and
sex ratios (Kuo & Janzen 2003, 2004). I conducted two simulations using current estimates of
population size and sex ratios from mark-recapture data collected for E. blandingii, C. picta, and
92
C. serpentina at the Will 3 site (see Fig. 3.3, Banning et al. 2006). Demographic parameters
remained constant for the 300-year duration. Estimates of longevity and ages of maturity for
each species were obtained from estimates reported from long-term studies and datasets
(Congdon et al. 1993, Congdon et al. 1994, Congdon et al. 2003, McGuire 2011). For the first
simulation I selected the random mating system option, and for the second simulation I selected a
skewed mating system option (i.e. one male sires all offspring each year) to model potential
effects of demographic stochasticity.
RESULTS
Amplification success, linkage disequilibrium, and Hardy-Weinberg equilibrium
I successfully genotyped 110 adult E. blandingii for 14 of the 15 microsatellite loci
(Table 3.1). One individual was only genotyped for 12 loci but was included in analyses. One
locus (GmuD90) could not be confidently scored because of inconsistent amplification and was
excluded from analyses. Presence of homozygous excess was detected at the Will 1 site for
GmuA18 and at Grundy site for GmuD70. Nevertheless, the occurrence of null alleles at these
loci is unlikely because tests of known mother-offspring genotype comparisons during parentage
analyses failed to produce any genotype mismatches (i.e. indication of null alleles; see Chapter
Five). Deviations from HWE were not detected after sequential Bonferroni correction (Table
3.2). Significant linkage disequilibrium (P = 0.0004; adjusted = 0.0006) was detected between
GmuD121 and GmuD21 for the Grundy site after Bonferroni correction but these loci were
retained for analyses because the significant relationship was restricted to one site.
93
Chrysemys picta
I successfully genotyped 331 adult C. picta for eight of the 15 microsatellite loci (Table
3.1). Eighteen individuals were genotyped for 6-7 loci and were included in analyses. One of the
successful loci (Eb17) was fixed for the Grundy County site but polymorphic for the Will
County sites. The seven remaining loci were excluded from analyses for various reasons:
GmuD90 and Eb19 could not be confidently scored because of inconsistent amplification,
GmuD121, GmuD87, and GmuA18 appeared to have null alleles (i.e., many samples failed to
amplify, homozygous excess), and Eb09 and BATC9 each only exhibited two alleles one repeat
motif apart and could not be confidently scored because of stutter patterns. Deviations from
HWE and significant linkage disequilibrium were not detected after sequential Bonferroni
correction (Table 3.2).
Chelydra serpentina
I successfully genotyped 83 adult C. serpentina for six of the seven microsatellite loci
(Table 3.1). One individual was only genotyped for two loci but was included in analyses. One
of the six successful loci (MteD2) was fixed for all study sites and was subsequently removed
from further analyses. The seventh locus (MteD106) could not be confidently scored and was
also excluded from analyses. The Grundy site was fixed and the Will 1 site only had one
heterozygote for the MteC1 locus; however, because this locus exhibited low polymorphism, the
lack of heterozygotes in the two sites was likely just an artifact of small sample size. Presence of
homozygous excess was detected at the Will 1 site for MteD9 but this also could be attributed to
small sample size. All loci conformed to the assumptions of HWE (Table 3.2). Significant
linkage disequilibrium (P = 0.002; adjusted = 0.005) was detected between MteC1 and
94
MteC112 for the Will 2 site after Bonferroni correction, but linkage comparisons across all sites
were not significant.
Genetic diversity within species across sites

For the 14 successful loci, I identified two to 13 alleles at each locus across all study sites
(Table 3.2; Appendix I). Interestingly, one individual from Will 3 was genotyped for three alleles
at three different loci (BATC9, GmuD70, and GmuD87) in each of two independent samples that
were collected in different years. I included this individual in subsequent analyses, but for the
triploid loci I only retained two of the three alleles that were most frequently observed in the
Will 3 site. Allelic richness and private allele richness were estimated from 22 gene copies in
each site to account for sample size variation. Mean number of observed alleles was greatest in
the intact site (Grundy) but measures of allelic richness were similar between the Grundy and
Will 1 sites (Table 3.2). Mean observed and expected heterozygosity were similar for all sites
and mean inbreeding coefficients did not indicate a loss of genetic diversity (Table 3.2).
Comparisons of genetic diversity (AR, PAR, Ho) did not differ between the intact Grundy County
and the fragmented Will County sites (Wilcoxon tests; P > 0.074, adjusted = 0.017).
Chrysemys picta
For the eight successful loci, I identified two to 73 alleles at each locus across all study
sites (Table 3.2; Appendix I). Allelic richness and private allele richness were estimated from 89
gene copies in each site to account for sample size variation. Mean allelic richness and total
number of private alleles were greatest for the fragmented Will County sites (Table 3.2).
95
However, mean observed heterozygosity and mean expected heterozygosity were similar among
sites (Table 3.2). Comparisons of genetic diversity (AR, PAR, Ho) did not differ between the intact
Grundy County and the fragmented Will County sites (Wilcoxon tests; P > 0.093, adjusted =
0.017).
Chelydra serpentina
For the five successful loci, I identified two to 16 alleles at each locus across all study
sites (Table 3.2; Appendix I). Allelic richness and private allele richness were estimated from 20
gene copies in each site to account for sample size variation. Mean allelic richness and total
number of private alleles were greatest for the Will 3 and Grundy County sites (Table 3.2). Mean
observed heterozygosity and mean expected heterozygosity varied slightly among sites and were
highest for the Will 2 site (Table 3.2). Comparisons of genetic diversity (AR, PAR, Ho) did not
differ between the intact Grundy County and the fragmented Will County sites (Wilcoxon tests;
P > 0.128, adjusted = 0.017).
Genetic divergence within species

For E. blandingii, pairwise FST analysis detected significant divergence between the
Grundy County and each of the Will County sites and between the Will 1 and Will 3 sites before
and after sequential Bonferroni correction (Table 3.3). For C. picta, significant divergence was
detected between Will 1 and Will 3 sites before but not after Bonferroni correction (Table 3.3).
For C. serpentina, divergence was detected between the Will 1 and Will 2 and between the Will
2 and Will 3 sites before and after Bonferroni correction (Table 3.3). AMOVA indicated weak
96
but significant structure among sites for E. blandingii (FST = 0.020, P = 0.001), C. picta (FST =
0.002, P = 0.010), and C. serpentina (FST = 0.011, P = 0.010).
Both simulations (with and without prior location information) of the Bayesian clustering
method indicated that there were three optimal clusters for E. blandingii and two optimal clusters
for C. picta and C. serpentina. However, the program failed to consistently assign individuals to
their respective sampling locations and assigned large proportions of individuals from one
location to more than one cluster indicating a lack of strong genetic divergence among sites (Fig.
3.2A-C).
Genetic divergence among species

For divergence among species, mean values of Dest and GST were low (< 0.04) and
patterns of divergence were inconsistent between C. picta and C. serpentina (Table 3.4). For,
Dest, C. serpentina was the least divergent among sites (CI included zero) but for GST, C.
serpentina was as highly divergent as E. blandingii. Further, significant differences in
divergence (95 CIs did not overlap) were only detected in GST comparisons; E. blandingii and C.
serpentina were more divergent across sites than C. picta.
Emydoidea blandingii exhibited subtle patterns of male-biased gene flow across all sites
but these patterns were more pronounced across fragmented sites (Table 3.5). Only FIS values
across fragmented sites were significantly larger in males than females (Table 3.5). No
significant differences in sex-biased dispersal were detected for C. picta or C. serpentina. In C.
picta, subtle patterns of male-biased gene flow were evident for mAIc and vAIc values but not
97
FST (Table 3.5). Further, FIS values were greater for female C. picta in both scenarios and little
difference between values was observed between fragmented sites only and all sites. In C.
serpentina, both scenarios showed subtle mixed patterns of sex-biased gene flow (Table 3.5). For
males, only FIS values indicated male-biased gene flow; whereas for females, vAIc and FST
indicated female-biased gene flow. Further, mAIc values (i.e. positive and negative) switched
between males and females in the comparison between fragmented sites only and all sites (Table
3.5).
Bottlenecks
No evidence of a past bottleneck (significant heterozygosity excess) was detected in any
of the fragmented Will County sites or the intact Grundy County site for E. blandingii (P = 0.770.96), C. picta (P = 0.95-1.00), or C. serpentina (P = 0.44-0.97) regardless of TPM mutation
parameters. All species also maintained a normal L-shaped distribution of allele frequencies
across sites, indicating no substantial loss of rare alleles that would be expected in a bottlenecked
population. The M-ratio tests also failed to show evidence of population declines (M > Mc) in all
species across all sites.

In both simulations of future genetic drift based on current demographic parameters and
allele frequencies, observed number of alleles decreased more quickly than observed
heterozygosity over the 300 year period (Fig. 3.3). Overall, loss of genetic diversity was most
pronounced in E. blandingii compared to the other two species. For the random mating
simulation, 88%, 97%, and 99% of heterozygosity was retained and 72%, 95%, and 94% of
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allelic diversity was retained for E. blandingii, C. serpentina, and C. picta, respectively after 300
years (Fig. 3.3). For the skewed mating simulation, resulting levels of genetic diversity were
lower compared to those of the random mating system but patterns of loss between the two
simulations varied among species. For example, patterns of genetic drift in E. blandingii were
similar regardless of mating system but C. serpentina and C. picta lost more heterozygosity (3%
and 4%) and substantially more allelic diversity (9% and 19%) in the skewed mating system
compared to the random mating system.
DISCUSSION
Overall, within the Lower Des Plaines River Valley (LDPRV) I found little evidence that
E. blandingii, C. picta, and C. serpentina in fragmented sites had less genetic variation when
compared to those in an intact site. All species demonstrated moderate to high levels of genetic
diversity. Further, I detected little genetic divergence among sites; however FST values among
sites varied by species. Gene flow was male-biased in E. blandingii across the fragmented sites
but differences in dispersal between males and females in C. picta and C. serpentina were not
strong. I found no evidence of genetic population bottlenecks in any species but simulations of
future genetic diversity suggest that E. blandingii is more vulnerable to loss of genetic diversity
than C. picta or C. serpentina.
Levels of genetic diversity

Comparisons of within-species levels genetic diversity observed across the LDPRV sites
were lower in E. blandingii when qualitatively compared to C. picta and C. serpentina.
However, estimates for all species were moderate and comparable to levels reported in other
99
freshwater turtles (Kuo & Janzen 2004,Tessier et al. 2005,Pearse et al. 2006, Castellano et al.
2009, Escalona et al. 2009, Ye et al. 2009, Spradling et al. 2010, Molnr et al. 2011). In my
study, inbreeding coefficients did not indicate inbreeding within species at any site. Estimates of
observed heterozygosity in previous E. blandingii studies that sampled 10 individuals/site
ranged from 0.71-0.80 in Illinois (Mockford et al. 2007, Klut 2011) and 0.61-0.64 in other
Midwest populations (Mockford et al. 2007). Differences in levels of genetic diversity among
species and studies can be attributed variability in locus polymorphism (Rubinsztein et al. 1995)
as well as the number of loci used to estimate diversity parameters.
This is one of the first studies known to report population genetic structure and gene flow
for C. picta and C. serpentina. Both of these species are common throughout their respective
geographic distributions but have received less attention than species of conservation concern
such as E. blandingii. With the exception of a DNA fingerprinting study that examined the
genetic diversity of C. picta between small and large wetland sites (Parker & Whiteman 1993),
previous genetic studies of C. picta and C. serpentina have focused on parentage analysis,
genetic mating systems, and assessments of multiple paternity (Galbraith 1993, Pearse & Avise
2001, Pearse et al. 2001, 2002, McGuire 2011, McGuire et al. 2011) and taxonomic relationships
(Phillips et al. 1996, Starkey et al. 2003).
Because turtle species examined in my study vary considerably in life history traits
(Congdon et al. 1993, Congdon et al. 1994, Congdon et al. 2003, McGuire 2011, McGuire et al.
2011), spatial ecology (Chapter One), and habitat use (Chapter Two), I had expected to find
differences in patterns of genetic diversity among species between the intact and fragmented
populations. Specifically, I had predicted fragmented E. blandingii populations to have lost more
genetic diversity and be more divergent between fragmented sites and the intact site than the two
100
common species, C. picta and C. serpentina. In a similar study that used DNA fingerprinting,
Parker & Whiteman (1993) found that the rare spotted turtle (Clemmys guttata) exhibited greater
differences in genetic diversity between small and large wetland complexes compared to the
abundant C. picta. However, I failed to detect significant differences in genetic diversity for any
of the three species between sites. Each of these species is capable of long-distance movements
via the Des Plaines River (Chapter One); thus, vagility coupled with long generation times
(Avise et al. 1992) and relatively recent fragmentation (Bennett et al. 2010) could account for
the lag in detectable loss of genetic diversity in fragmented sites.
Measures of genetic divergence

I detected significant pairwise FST divergence in E. blandingii and C. serpentina.
Although FST values were low, E. blandingii was divergent between the intact and each of
fragmented sites as well as between two of the fragmented sites (Will 1 and Will 3). Conversely,
C. serpentina was only divergent between Will 2 and Will 1, as well as Will 2 and Will 3. I
suspect that the levels of divergence in C. serpentina are attributed to variation in sample size.
Samples from female C. serpentina are lacking from the Will 1 and Will 3 sites compared to
Will 2 and considering that female C. serpentina are known to be philopatric to nesting sites
(Congdon et al. 1987), a male-biased sample pool could impact levels of divergence among sites.
In the direct comparisons among species, E. blandingii was the most divergent for both
pairwise estimates (Dest and GST). However, patterns of divergence were not consistent for C.
picta and C. serpentina. Further, significant differences among species were only detected using
the GST estimates; E. blandingii and C. serpentina were more divergent than C. picta. The
discrepancies between these two measures may be attributed to differences in the underlying
101
dependencies in heterozygosity and mutation rates (Hedrick 2005, Jost 2008, Ryman & Leimar
2009). Further, accuracy of GST in measuring differentiation has been criticized (Jost 2008,
Gerlach et al. 2010) and thus should be interpreted with caution. Variation in life history traits
(e.g. longer generation time) could explain why E. blandingii is more divergent among sites
compared to the other two species.
Females of many species of turtles are philopatric to nesting locations, including E.
blandingii (Congdon et al. 1983), C. picta (Valenzuela & Janzen 2001, Rowe et al. 2005), and C.
serpentina (Congdon et al. 1987), whereas males are considered to be the dispersing sex (but see
Sheridan et al. 2010). However, in this study sex biased gene flow was only evident for E.
blandingii and was more apparent in fragmented sites alone than when including the intact
Grundy County site. If the dispersing sex is more genetically similar across sites than the
philopatric sex and contemporary fragmentation prevents successful dispersal among sites, then
FIS values should increase in the dispersing sex (i.e. males). Assemblages of E. blandingii found
in the LDPRV fragmented sites are small and biased towards females (Banning 2006, Banning et
al. 2006, Dreslik et al. 2011). Thus, fewer numbers of males across the fragmented sites could
explain the stronger bias in levels of male FIS compared to the scenario that included the intact
site. Alternatively, stronger evidence for male-biased gene flow in the fragmented sites may be
related to their closer proximity and potential for higher levels of historical gene flow than the
more distant intact site. The lack of sex-biased gene flow in C. picta and C. serpentina could be
caused by either a lack of male dispersers or a combination of male and female dispersers. For C.
picta, high genetic diversity and no differentiation across sites suggest that gene flow was
102
historically high across sites and lends support to the latter dispersal explanation. Female natal
philopatry as well as male and female dispersal has been reported in the diamondback terrapin
(Malaclemys terrapin, Sheridan et al. 2010). For C. serpentina, because evidence for
differentiation across sites is unclear, sex-biased gene flow may be present but undetected.
Bottlenecks
Although suitable turtle habitat has been lost and fragmented within the LDPRV,
evidence of recent population declines was not evident for any species. Lack of genetic
divergence and population bottlenecks, even in small isolated sites, are not uncommon in turtles
(Parker & Whiteman 1993, Rubin et al. 2001a, Kuo & Janzen 2004, Mockford et al. 2007,
Bennett et al. 2010, Spradling et al. 2010, Klut 2011) and have been attributed to a combination
of long generation times, low metabolic and mutation rates (Avise et al. 1992), and relatively
recent anthropogenic habitat fragmentation (Bennett et al. 2010). Both, spatial and temporal
scale can affect power to detect patterns in landscape genetic studies and a lag time can exist
between landscape change and a response in biological processes (Anderson et al. 2010). The
turtle gene pools sampled in my study occur within a relatively localized scale; a 50 km stretch
of the LDPRV. Historically, these groups were likely panmictic and movement and gene flow
could occur throughout matrices of prairie and wetland habitats without anthropogenic
impediment. Although contemporary movement among these remnant populations has been
restricted to dispersal via the Des Plaines River and subtle differentiation is evident only in E.
blandingii across sites, not enough time (i.e. generations) may have yet passed to detect the
subsequent loss of genetic diversity and gene flow in C. picta and C. serpentina.
103

Simulations of future loss of genetic diversity demonstrated that differences in speciesspecific traits such as age of maturity, longevity, sex ratio, and abundance appear to affect the
rates of genetic drift among E. blandingii, C. picta, and C. serpentina within the LDPRV. Loss
of genetic diversity was substantially higher in E. blandingii than for C. picta or C. serpentina.
This can be explained by the long time to maturity, greater longevity, and drastically smaller
estimated population size in E. blandingii compared to C. picta and C. serpentina. Although C.
serpentina appear to be more stable compared to E. blandingii, simulations of future genetic
diversity suggest that C. serpentina is more vulnerable to genetic loss than C. picta. Its
intermediate position of conservation concern is likely a result of the combination of
demographic parameters and ecological specialization of C. serpentina. On one hand, this
species is relatively abundant (Banning et al. 2006, Dreslik et al. 2011), capable of long-distance
aquatic movements (Chapter One) that can potentially maintain gene flow among populations
and is a habitat generalist that readily uses poorer quality habitats including the Des Plaines
River (Chapter Two). However, C. serpentina also exhibits a longer time to sexual maturity and
a longer life span than C. picta that is more similar to E. blandingii in these regards (Congdon et
al. 1993, 1994, 2003). Alterations in the mating system settings (random vs. skewed) had the
greatest impact on C. serpentina and C. picta. However, skewed mating extremes (i.e. only one
male siring all offspring) do not reflect actual mating systems reported in populations of C.
serpentina and C. picta (Pearse and Avise 2001, Pearse et al. 2002, McGuire 2011) and are
unlikely for population with large numbers of individuals as estimated for the Will 3 site.
104
Conservation Implications
Loss of genetic diversity and divergence in fragmented sites compared to an intact site
was not apparent within the LDPRV. However, lack of contemporary dispersal (Chapter One)
and gene flow (Chapter Four) between sites is potentially masked by long-generation times and
relatively recent landscape fragmentation. Long-term loss of genetic diversity is possible in all
three turtle species but is particularly imminent in E. blandingii because of lower abundance and
longer generation time of this species across sites compared to C. picta and C. serpentina.
Because populations do not appear to be substantially different genetically, long-term
management of LDPRV sites should try to maintain some level of gene flow and consider
actions such as translocation of head-started hatchlings.
105
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Ecology, 12, 11-20.
Ye R, Zheng R, Wang L, Du W (2009) Polymorphic microsatellite loci in the Chinese pond
turtle (Chinemys reevesii). Conservation Genetics 10, 1045-1048.
115
TABLES
116
Table 3.1 Numbers of male (), female (), unknown (U), and total (T) adult individuals
successfully amplified across selected microsatellite loci in E. blandingii, C. picta, and C.
serpentina from three fragmented sites (Will 1-3) and one intact site (Grundy) within the Lower
Des Plaines River Valley.
E. blandingii
C. picta
C. serpentina
Site
Will 1
Will 2
Will 3
Grundy
8
3
10
23
14
8
20
24
22
11
30
47
68
45
57
30
38
26
53
14
106
71
110
44
15
17
12
8
5
12
5
2
1
6
0
0
21
35
17
10
Total
44
66
110
200
131
331
52
24
83
117
Table 3.2 Mean estimates for number of alleles (#A), allelic richness (AR), private allele richness
(PAR), observed heterozygosity (Ho), expected heterozygosity (He), inbreeding coefficients (FIS),
and probability of Hardy-Weinberg deviation (PHWE) for E. blandingii, C. picta, and C.
serpentina sampled from three fragmented (Will 1-3) and one intact site (Grundy) within the
Lower Des Plaines River Valley. Data were derived from microsatellite analysis with numbers of
loci analyzed in E. blandingii = 14, C. picta = 8, and C. serpentina = 5. Sample sizes for each
species at each site are provided in Table 3.1.
Site
AR
PAR
Ho
He
FIS
PHWE
4.9
4.1
4.9
5.6
4.2
4.1
4.0
4.3
0.42
0.38
0.17
0.43
0.535
0.552
0.552
0.506
0.532
0.535
0.537
0.522
0.034
-0.014
-0.029
0.001
0.48
0.53
0.67
0.56
15.0
14.9
14.4
11.5
11.8
12.7
11.9
11.4
0.91
1.40
0.93
0.53
0.662
0.664
0.662
0.627
0.660
0.652
0.659
0.639
-0.011
-0.014
-0.020
0.002
0.66
0.65
0.57
0.58
6.2
7.2
6.8
5.8
5.1
5.4
5.8
5.8
0.05
0.21
0.56
0.25
0.543
0.691
0.576
0.620
0.579
0.651
0.578
0.619
0.024
-0.058
-0.006
-0.003
0.46
0.65
0.84
0.39
E. blandingii
Will 1
Will 2
Will 3
Grundy
C. picta
Will 1
Will 2
Will 3
Grundy
C. serpentina
Will 1
Will 2
Will 3
Grundy
118
Table 3.3 Pairwise estimates of FST (below diagonal) and p-values estimated from 999
permutations (above diagonal) for A) E. blandingii, B) C. picta, and C) C. serpentina among
three fragmented (Will 1-3) and one intact site (Grundy) within the Lower Des Plaines River
Valley. Significant FST values after sequential Bonferroni correction are denoted with an *.
Genetic data were derived from microsatellite DNA analysis with numbers of loci analyzed in E.
blandingii = 14, C. picta = 8, and C. serpentina = 5. Sample sizes for each species at each site
are shown in Table 3.1.
A) E. blandingii
Will 1
Will 2
Will 3
Grundy
Will 1
Will 2
Will 3
Grundy
---0.014
0.023*
0.018*
0.072
---0.008
0.029*
0.003
0.169
---0.026*
0.002
0.002
0.001
----
Will 1
Will 2
Will 3
Grundy
---0.001
0.003
0.002
0.155
---0.001
0.003
0.011
0.155
---0.003
0.134
0.083
0.081
----
Will 1
Will 2
Will 3
Grundy
---0.017*
0.000
0.000
0.010
---0.037*
0.000
0.475
0.001
---0.011
0.408
0.421
0.167
----
B) C. picta
Will 1
Will 2
Will 3
Grundy
C) C. serpentina
Will 1
Will 2
Will 3
Grundy
119
Table 3.4 Standardized estimates of Dest and GST (with 95% Confidence Intervals in parentheses)
for E. blandingii, C. picta, and C. serpentina sampled from three fragmented (Will 1-3) and one
intact site (Grundy) within the Lower Des Plaines River Valley. Genetic data were derived from
microsatellite DNA analysis with numbers of loci analyzed in E. blandingii = 14, C. picta = 8,
and C. serpentina = 5. Sample sizes for each species at each site are shown in Table 3.1.
Dest (95% CIs)
GST (95% CIs)
E. blandingii
0.039 (0.018-0.063)
0.031 (0.020-0.042)
C. picta
0.017 (0.003-0.032)
0.008 (0.005-0.012)
C. serpentina
0.009 (-0.039-0.065)
0.031 (0.026-0.051)
Species
120
0.012 -0.027
-0.006 0.014
C. serpentina
All Sites
Frag. Sites
0.303
0.215
-0.200
-0.150
C. picta
All Sites
Frag. Sites
0.050
0.173
-0.072
-0.347
E. blandingii
All Sites
Frag. Sites
Sites
mAIc
0.54
0.49
0.07
0.14
0.46
0.34
2.9
3.1
9.7
9.9
19.5
19.0
4.2
4.1
8.6
9.0
16.6
14.4
vAIc
0.82
0.76
0.19
0.27
0.41
0.24
0.005
0.002
0.023
0.025
0.012 -0.022
0.022 -0.014
0.001
0.001
0.026
0.009
FST
0.96
0.97
0.11
0.36
0.58
0.32
0.017
0.012
0.024 0.005
0.016 -0.001
-0.008
-0.011
0.053 -0.006
0.095 -0.045
FIS
0.57
0.5
0.91
0.89
121
0.08
0.004*
Table 3.5 Tests for differences in mean assignment indices (mAIc), variance in assignment indices (vAIc), FST, and FIS between male
and female E. blandingii, C. picta, and C. serpentina from sites within the Lower Des Plaines River Valley. Parameters were estimated
sites using the biased dispersal option in FSTAT V. 2.9.3.2 (Goudet 1995). Significance is indicated by an * and = 0.05.
FIGURES
122
Fig. 3.1 Location of turtle sampling sites in northeastern Illinois, USA. Sites Will 1, Will 2, Will
3, and Grundy are indicated by red stars and are located from north to south, respectively, along
the Des Plaines River.
DU PAGE
Chicago
JO DAVIESS
STEPHENSON
WINNEBAGO
BOONE
MCHENRY
LAKE
KENDALL
CARROLL
WILL
OGLE
DE KALB
KANE
COOK
DU PAGE
WHITESIDE
LEE
WILL
KENDALL
LA SALLE
HENRY
IS
BUREAU
ND
LA
CK
RO
GRUNDY
MERCER
PUTNAM
KANKAKEE
KNOX
MARSHALL
ST ARK
WARREN
LIVINGSTON
PE ORIA
O
RS
DE
N
HE
IROQUOIS
N
WOODFORD
GRUNDY
FULTON
MC LEAN
MCDONOUGH
HANCOCK
FORD
TAZEWELL
VERMILION
CHAMPAIGN
MASO N
SCHUYLER
LOGAN
DE WITT
ADAMS
BR OWN
MENARD
PIATT
CA SS
MACON
MORGAN
SANGAMON
PIKE
DO UGLAS
SCOTT
EDGAR
MOUL TRIE
CHRISTIAN
SHELBY
GREENE
COLES
MACOUPIN
CLAR K
CALH
CUMBERLAND
MON TGOMERY
OUN
KANKAKEE
FAYETTE
JERSEY
EFFINGHAM
MADISON
CR AWFORD
JASPER
BOND
CLAY
RICHLAND
MARION
LAWRENCE
CLINTON
ST. CLAIR
DS
ED
WAR
JEFFERSON
WAB
WASHINGTON
AS
H
WAYNE
MON ROE
RANDOLPH
HAMILTON
PERRY
WHITE
FRANKLIN
JACKSON
SALINE
GALLATIN
WILL IAMSON
UNION
JOHNSON
POPE
HAR DIN
0 10 20
40 Kilometers
AL
MASSAC
EX
PULASKI
AN
DE
R
123
Fig. 3.2 Bayesian clustering results based on the LOCPRIOR option in STRUCTURE 2.3.3
(Pritchard et al. 2000) A) E. blandingii (3 clusters), B) C. picta (2 clusters), and C) C. serpentina
(2 clusters) among three fragmented (Will 1-3) and one intact site (Grundy) within the Lower
Des Plaines River Valley. DNA analysis with numbers of loci analyzed in E. blandingii = 14, C.
picta = 8, and C. serpentina = 5. Sample sizes for each species at each site are shown in Table
3.1.
A)
B)
C)
124
Fig. 3.3 Comparison of predicted genetic variation retained for observed # alleles (OA) and
observed heterozygosity (Ho) over 300 years in populations of three turtle species from a
preserve in Will County, Illinois. Simulations included the below demographic estimates and life
history traits and were performed for random and skewed mating systems using the program
BOTTLESIM v2.6 (Kuo and Janzen 2003).
Ho Random
H
Ho Skewed
OOA Random
OA Skewed
100
95
90
% 85
Gen 80
etic 75
Vari 70
atio 65
n 100
Reta 95
ined 90
E. blandingii
Populations Size: 43
Sex Ratio: 1M:2F
Longevity: 75 yrs
Age Maturity: 14
yrs
85
C. picta
Sex Ratio: 1.3M:1F
Longevity: 50 yrs
Age Maturity: 7 yrs
80
75
70
65
100
95
90
85
C. serpentina
Sex Ratio: 1M:1.2F
Longevity: 55 yrs
Age Maturity: 11 yrs
80
75
70
65
0
50
100
150
200
250
300
Years Simulated
125
CHAPTER 4
MATING SYSTEM AND REPRODUCTIVE SUCCESS IN A FRAGMENTED
POPULATION OF BLANDINGS TURTLES (EMYDOIDEA BLANDINGII)
INTRODUCTION
Data on reproductive ecology are important for understanding population dynamics and
demographics, and are an integral part of conservation planning. For example, the mating system
of a species is an important life history component because the number of reproducing
individuals directly influences effective population size Ne, genetic drift, and inbreeding
(Caballero 1994) which have conservation implications for small fragmented populations (Willi
et al. 2006, Allendorf and Luikart 2007, Mills 2007). In addition, fragmentation can alter
distribution of mates (Lane et al. 2011) and elevate inbreeding risk (Andersen et al. 2004, Banks
et al. 2005) by confounding dispersal patterns and disrupting gene flow among populations
(Moore et al. 2008a). Remnant populations with limited dispersal may consist predominantly of
related individuals and have lowered reproductive success because of inbreeding depression
(Mills and Smouse 1994). In addition, because of Allee effects, small populations may simply
lack sufficient numbers of individuals to ensure adequate encounter rates to facilitate mating
(Tainaka and Itoh 1996, Stephens and Sunderland 1999, Dale 2001, Robertson and Butler 2009).
A mating system consists of number of mates, method of acquiring mates, pair bond
characteristics, and manner of parental care (Emlen and Oring 1977). Compared to other taxa,
non-avian reptiles have less complex mating systems because they typically lack pair bonds and
parental care (Pearse and Avise 2001, Uller and Olsson 2008). However, widespread
reproductive strategies among reptile taxa such as multiple paternity and sperm storage are not
straightforward and lead to misinterpretations of mating system patterns (Uller and Olsson
126
2008). For example, single clutches sired by multiple males could stem from fertilization from
stored and recently inseminated sperm (Pearse et al. 2001, 2002) and confound inferences of
polyandry versus seasonal monogamy (Uller and Olsson 2008). Anecdotal observations of
courtship and mating attempts are uncommon (Sexton 1959a) and do not necessarily equate to
reproductive success (Fitzsimmons 1998). Further, mechanisms of cryptic female choice and
sperm competition are not well documented (Galbraith 1993, Eberhard 1996, Uller and Olsson
2008, Olsson et al. 2010). Therefore, studies of mating systems are most informative when
behavioral observations can be paired with parentage analysis in naturally occurring populations
(Pearse et al. 2002, Moore et al. 2009, Uller and Olsson 2008).
In this study, I assessed the mating system and reproductive success of two adjacent
populations of Blandings turtle (Emydoidea blandingii) within a fragmented landscape by
pairing field observations of mating behavior with genetic parentage analysis of offspring.
Emydoidea blandingii is a species of conservation concern throughout its range largely because
of habitat loss (Ernst and Lovich 2009). This species is capable of long overland movements (up
to > 1 km) during nesting forays (Sexton 1995, Piepgras and Lang 2000, Joyal et al. 2001) and
between wetlands (Piepgras and Lang 2000, Rowe and Moll 1991) but anthropogenic barriers
such as roads prevent successful dispersal and gene flow in turtle species (Gibbs and Steen
2005). Recent studies of the mating system and parentage in E. blandingii have demonstrated
multiple paternity (15-81%), repeat paternity (69%), and possible sperm storage (Refsnider 2009,
McGuire 2011), but no study to date has compared behavioral observations of courtship to
parentage or assessed the mating system of this species in fragmented landscapes.
My objectives were to determine 1) timing and frequency of mating attempts, 2) number

of potential mates among individuals, 3) number of offspring and clutches sired by males across
127
four sequential breeding seasons, 4) determinants of male reproductive success, and 5) effects of
male-female pair relatedness on reproductive success. Specifically, I predicted observations of
mating attempts would correspond with DNA parentage analysis. Because larger males have
larger home ranges (Chapter One), I expected that males with larger home range sizes should
encounter more females and have more successful matings, which would lead to higher
reproductive success. Body size has been shown to be a predictor of greater reproductive success
in reptiles (Moore et al. 2009, Olsson et al. 2010, Tuberville et al. 2011), thus I also predicted
that larger males could also sire more offspring because they would be more likely to defend
females from other males and coerce females into successful matings. Because matings between
related individuals (i.e. inbreeding) can reduce reproductive success, I expected related malefemale pairs to have lower mating and hatching success than un-related pairs. I also expected low
levels of gene flow because few individuals have been observed to move between sites (Chapter
One).
METHODS
Reproductive ecology of Emydoidea blandingii
Emydoidea blandingii are long-lived (70+ years) with delayed sexual maturity at 14-20
years of age (Congdon et al. 1993). Females ovulate in May (Gibbons 1968) but the reproductive
cycle is unknown in male E. blandingii (Ernst and Lovich 2009). Anecdotal accounts of
courtship in natural populations of E. blandingii have been observed throughout the year (Carr
1952, Graham and Doyle 1979), and suggest that the timing of mating and fertilization may be
decoupled (Devine 1984, Uller and Olsson 2008). Mature females produce one clutch per year
but may not reproduce every year (Congdon et al. 1983, Congdon and van Loben Sels, 1993,
128
Banning 2007) and in Michigan, among-year clutch frequency and multiple paternity of clutches
increases with female age (Congdon and van Loben Sels 1993, McGuire 2011). In Will County,
Illinois, clutch size of E. blandingii may range from 8-19 eggs but average clutch size is 11-13
eggs (Banning 2007, Dreslik et al. 2011). Nesting occurs in evenings during late-May-early July
with females being philopatric to nesting areas (Congdon et al. 1983, Banning 2007). Hatchlings
emerge in August October (Congdon et al. 1983, Anthonysamy et al. unpublished data).
Field methods
I radio-tracked adult E. blandingii 3-7 times per week during a radio-telemetry study
conducted from 2006-2009 at two semi-connected preserves (Will 1 and Will 2) in Will County,
Illinois. Radio-transmitters were removed from males in October 2009 but transmitters were left
on females until June 2010 to collect an additional year of nesting data. Details on study sites and
radio-telemetry methods are described in Chapter One. During radio-telemetry, I recorded all
observations of mating attempts (mounting of females by males). Observations of mounting were
considered to be only mating attempts as this behavior is part of a sequence of courtship that
precedes copulation in E. blandingii but does not necessarily indicate that successful copulation
occurred (Baker and Gillingham 1983). Repeated mating attempts between the same pairs were
considered to be distinct events if they were separated by > 3 days (Rovero et al. 1999). I
recorded temporal patterns of mating attempts, number of mating attempts, and number of
potential mates observed for each turtle.
I determined whether females were gravid by palpating the inguinal pocket for presence
of eggs during nesting season (late Mayearly July) and by radiography (Gibbons and Greene
1979). I obtained clutches during 2007-2010 by 1) radio-tracking females to nesting locations,
129
protecting nests with predator exclusion cages, and harvesting hatchlings from the cages; 2)
radio-tracking females to nesting locations and collecting eggs from nest chambers; or 3)
transporting gravid females to the Willowbrook Wildlife Center (Glen Ellyn, Illinois) to induce
egg-laying with intramuscular injections of oxytocin (7.5 units/kg) or a combination of oxytocin
(1.5 units/kg) and lutalyse (1.5 mg/kg). Clutches obtained from nest chambers were either placed
in an incubator at a temperature of approximately 27-30 C or were incubated at room
temperature in plastic shoeboxes filled with moistened vermiculite. Clutches from induced
females were placed in incubators with moistened vermiculite (constant temperature of 28 C) at
the Willowbrook Wildlife Center. Nests and incubated clutches were monitored periodically
until hatchling emergence to determine hatching success and to collect tissue samples. I
calculated hatching success as the proportion of live, non-deformed, and active hatchlings
sampled from the total number of eggs collected for a clutch.
Lab methods
I collected blood (0.1-0.3 cc) from the sub-carapacial sinus (Fisher 2003) of adult turtles
using a 25 gauge needle and 1 cc syringe and collected 1-2 mm tail clips from hatchlings using
sterilized cuticle scissors. I stored tissues in 95% ethanol or Queens lysis buffer (Seutin et al.
1991) at -80C until DNA extraction. I extracted whole genomic DNA from tissue samples using
the QIAGEN DNeasy Blood & Tissue Kit (QIAGEN INC.) following the manufacturers protocol,
with the exception that I digested tissue samples overnight in a proteinase K solution.
For each DNA sample, I amplified 14 microsatellite loci using primers developed for E.
blandingii ([BTCA9; Libants et al. 2004] [Eb 09, Eb 17, Eb 19; Osentoski et al. 2002]) and bog
turtle (Glyptemys muhlenbergii; GmuD21, GmuD55, GmuD70, GmuD87, GmuD93, GmuD121,
130
GmuB08, GmuA18, GmuA19, GmuA32; King and Julian 2004). Polymerase chain reaction
(PCR) was carried out using the protocol described in Chapter Three. Fragment analysis of
resulting PCR products was carried out on an automated Applied Biosystems (ABI) Prism
3730xl sequencer at the W. M. Keck Center at the University of Illinois, Champaign. An internal
size standard was run with each sample (LIZ500). I scored alleles using GENEMAPPER 4.1
software (ABI) and identified possible null alleles, large allele dropout, and scoring errors due to
stutter peaks using MICRO-CHECKER 2.2.3 (van Oosterhout et al. 2004).
Genetic analyses
Using only adult samples, I tested for linkage disequilibrium (Markov Chain parameters:
10000 dememorisation steps, 500 batches, 5000 iterations) between all pairs of loci and tested
for departures from Hardy-Weinberg equilibrium for each locus using exact tests in GENEPOP 4.0
(Rousset 2008). I estimated allele frequencies, observed heterozygosity (Ho), and expected
heterozygosity (He) using GENALEX 6.41 (Peakall and Smouse 2006).
Paternity analysis
To determine the number of offspring and clutches sired among males, I assigned
paternity using GERUD 2.0 (Jones 2005) a software program that calculates exclusion
probabilities for loci and reconstructs parental genotypes from arrays of full or half-sib progeny.
Paternal genotypes were reconstructed based on genotypes of known mothers and their clutches
and were then compared to genotypes of the 11 sampled candidate males in the population. I also
used the results of the paternity analysis to determine if mating observations observed during
radio-telemetry corresponded to sired offspring.
131
Determinants of male reproductive success

I examined the Pearson-product moment correlation coefficient (r) between male traits
and reproductive success using the correlation (test.cor) function implemented in R software
2.13.2 (R Development Core Team 2011). Reproductive success was measured as total number
of offspring sired over the course of the study. Male traits included body size (carapace length,
CL), number of potential mates observed during radio-telemetry surveys, number of successful
mates inferred from paternity analysis, and home range size (HR). For home range, I used 95%
kernel density isopleth home range size that was estimated in Chapter One from all radiolocations collected for a respective male throughout its duration in the study. I also examined the
correlation between CL vs. number of potential mates, CL vs. number of successful mates, HR
vs. number of potential mates, and HR vs. number of successful mates. For all correlation
analyses except number of successful mates vs. number of offspring, CL vs. number of
successful mates, and CL vs. number of offspring I included only mature, radio-telemetered
males that had sufficient locations to estimate multi-year home range size (N=7). One additional
male with less than one year of location data was included in tests between number of successful
mates vs. offspring, CL vs. number of successful mates, and CL vs. number of offspring.
Genetic compatibility and reproductive success

Genetic compatibility or effect of inbreeding avoidance on number of successful mates
and clutch hatching success was assessed by estimating a relatedness coefficient (R) of malefemale pairs in the program ML-RELATE (Kalinowski et al. 2006). Relatedness coefficients range
from 0-1 with 0 indicating no relatedness and 1 indicating complete relatedness. I calculated a
Pearsons product-moment correlation coefficient (r) between relatedness coefficients (R) and
132
successful pairings and hatching success. Because hatching success was low in 2007 and 2008, I
tested the correlation between R and hatching success with and without the 2007 and 2008
data.
RESULTS
Reproductive and genetic data were collected from ten male and 19 female telemetry
subjects (Table 4.1). Genetic data were also collected from three additional males and one female
that were encountered during telemetry, but were not telemetry subjects. No other adult E.
blandingii were encountered during 131,444 hours of trapping at the Will 1 site; thus, the
numbers of males and females studied likely reflect a true female-biased sex ratio in the sites
(Dreslik et al. 2011).
Timing and frequency of mating behavior

I recorded 39 distinct mating attempts between eight adult males and 15 adult females
(Table 4.2). Mating attempts were observed throughout the active season except for June. Most
attempts occurred during the spring and fall; however the greatest number of monthly
observations was recorded in July (Fig.4.1). The total number of distinct mating attempts
observed per male ranged from 0-10 and the total number of potential mates observed with a
given male over the course the study ranged from 0-8 (Table 4.3). The total number of distinct
mating attempts per female ranged from 0-5 and the total number of potential mates observed
with a given female over the course the study ranged from 0-3. Two males (AXEL and RMEO)
were never observed engaging in mating attempts (Table 4.3) and so were excluded from further
analyses because although they exhibited secondary sexual characteristics, their size (plastron
133
lengths of 175 and 182 mm, respectively) suggests that they may not have been sexually mature
(Ernst and Lovich 2009). In addition no clutches were obtained from two females (FRAN and
CLET) because their transmitters failed early in the study and they were never recaptured. These
females were also subsequently excluded from further analyses to reduce bias in comparisons
between potential mating attempts and number of successful mates as inferred from paternity
analyses.
Number of potential mates

Mean number of potential mates observed during radio-telemetry surveys varied among
years and between telemetered males and females (Table 4.4A; Appendix J). Over four years of
radio telemetry, males and females on average were observed with 3.8 and 1.5 potential mates,
respectively. However, within-year estimates of potential mates were < 1 for two years in males
and each of the four years in females (not all turtles were observed in mating attempts in each
year). Average observed number of potential mates was lowest in 2006 (0.3) and highest in 2007
(1.3) but this variation likely reflects differences in the number of turtles radio-tracked during
each year or differences in detection ability among field researchers between years. Nevertheless,
males always averaged more potential mates than females (Table 4.4A). Four females were
never observed engaging in mating behavior but each produced at least one clutch during the
study. This demonstrates that we likely missed a number of mating observations during radiotelemetry surveys.
134
Clutch samples and hatching success

Most females ( 80%) produced a clutch in each nesting season in which they were
monitored. However, some females had incomplete reproductive histories because I failed to
locate six natural nests (four in 2007, two in 2008), transmitter failure before nesting season, or
extremely short duration of telemetry. Two natural nests were completely depredated and one
was partially depredated (clutch 27-08). One female (EDNA) monitored over four consecutive
nesting seasons (2007-2010) never produced a clutch.
I collected 35 clutches (whole and partial) from 16 females for a total of 272 hatchlings
with known mothers during 2007-2010 (Table 4.5). Number of hatchlings per clutch ranged from
1-18 hatchlings (mean = 8) and 28 clutches had at least three hatchlings; the minimum number
necessary to detect more than two paternal alleles in a clutch (i.e. multiple paternity). I obtained
hatchlings from one, two, and three clutches from five, seven, and four females, respectively,
which allowed me to examine repeat paternity among years (see Parentage and Quantification of
Mating Success section below).
Hatching success varied among years and among females but was lowest in 2007 and
2008 compared to 2009 and 2010 (Table 4.5) and is attributed to natural factors (i.e. flooding and
ant infestation in 2007) and possibly to extended handling of clutches (transport of eggs to field
station vs. immediate placement in incubator) and differences in incubation methods. Twenty-six
clutches from naturally and artificially incubated nests contained 1-16 unviable eggs with no
development, including four with a 100% failure rate (Table 4.5). Nine artificially incubated
clutches produced 1-3 egg-bound or malformed/lethargic hatchlings. I assumed that
malformed/lethargic individuals would not have emerged from the nest under natural
135
circumstances and did not consider them successful hatchlings when calculating hatching
success.
Genetic analyses
All 33 adults and 272 hatchlings (alive and dead) were successfully genotyped across 14
microsatellite loci, but one non-telemetered female and one hatchling only yielded genotypes for
12 loci. For sampled adults, all loci conformed to the assumptions of Hardy-Weinberg
equilibrium and no evidence of significant linkage disequilibrium was detected after Bonferroni
correction (Table 4.6). Because the turtles at the Will 1 and Will 2 preserves were once part of a
larger panmictic population and movement of individuals between sites has been documented
(Chapter 1), and because pairwise FST comparisons between Will 1 and Will 2 showed no
genetic differentiation (Chapter 3), I combined the two populations for subsequent analyses.
Male reproductive success

The program GERUD yielded either one (single paternity) or two (multiple paternity)
possible sires for each clutch tested. Combined exclusion probabilities for the 14 loci with one
parent known were > 0.99 (Table 4.6). Thus, paternity was easily established by comparing
reconstructed paternal genotypes to the genotypes of the candidate male samples. Paternity was
assigned to eight telemetered males and one un-sampled male (Table 4.5). The two telemetered
males without any mating observations (AXEL and RMEO) failed to sire any offspring. The total
number of offspring sired by each male was heavily skewed with one male siring 37% (N=102)
of hatchlings (Fig. 4.2) and 36% of clutches (N=11; Tables 4.3 & 4.5; Fig. 4.3).
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Quantification of mating success

Mean number of successful mating attempts varied among years and between telemetered
males and females (Table 4.4B; Appendix J). Over four consecutive breeding seasons (20072010), males and females averaged 2.6 and 1.3 different mates, respectively. Six males (JAY,
LIPA, EZRA, BIPA, VERN, and ZEB) sired offspring with 2-5 different females during the study.
Six females (VLMA, MAUD, MRTH, HART, EZMR, and SMLY) sired offspring with two different
males during the course of the study.
On 18 occasions, males attempted to mate with females with whom they were known to
sire offspring at some point during the study. Of the 39 courtship observations documented
during radio-telemetry, 22 observations could be compared to subsequent clutches acquired from
females (Table 4.2). The remaining observations could not be used for comparisons because two
females did not produce clutches during nesting seasons following observations that occurred in
the preceding fall or spring and for 15 observations, mating success could not be determined
because either nests could not be located, transmitter failure, or clutches had 100% hatch failure
with no embryonic development. When considering the 23 clutches that were collected during
the radio-telemetry project (clutches from 2007-2009 with at least one sampled hatchling) that
could have been linked with behavioral data, nine out of the 22 observations (41%) corresponded
with mating events that occurred during the fall or spring preceding the clutch (Tables 4.2 and
4.5). Conversely, there were 14 clutches with no apparent corresponding mating observation
during the preceding fall or spring; however five of those clutches could potentially have been
sired via stored sperm 1-2 years after the previous mating event but this is questionable because
in most of those cases the females produced clutches sired by different males during that 1-2 year
interim (Tables 4.2 and 4.5).
137

A significant positive correlation was observed for number of successful mates vs.
number of offspring (r = 0.78, df = 6, P = 0.022) but no other male trait (CL, HR, number of
potential mates) was related to number of offspring sired. No relationship between body size
(CL) and number of potential or successful mates and no relationship between HR and number
of potential or successful mates was detected.

Relatedness coefficients (R) calculated for observed male-female pairs during radiotelemetry and male-female pairs that sired clutches ranged from 0-0.52 (Tables 4.2 and 4.5). At
least 31% of observed pairings and 52% of clutches were produced from mated pairs with an R >
0. The correlation between relatedness and success of mating was negative but non-significant (r
= -0.18, df = 39, P = 0.272). Further, I found no indication that relatedness affected hatching
success with 2007 and 2008 clutches included (r = 0.09, df = 31, P = 0.631) or excluded (r =
0.17, df = 18, P = 0.461).
Detection of multiple paternity

Multiple paternity was detected in only three clutches (8%). Multiply sired clutches were
produced by three different females (Table 4.5). At least three different paternal alleles were
detected for the Eb09, BATC9, GmuD70, GmuA19, and GmuB08 loci in clutch 40-09, BATC9,
GmuD70, and GmuB08 in clutch 38-10, and GmuD70 in clutch 26-08. One male (BIPA)
contributed offspring in all three multiply sired clutches but sired a lower proportion of offspring
in two of those clutches (Table 4.5). Paternity assignments for some males and multiple paternity
138
in some clutches could have gone undetected because I failed to locate six natural nests and four
nests were completely unviable with no development and therefore no paternal DNA
contribution.
Evidence of sperm storage

Across season sperm storage was confirmed in one clutch (11-08) that was sired by a
male the nesting season (June 2008) following his death (August 2007). Circumstantial evidence
suggests other possible instances of sperm storage. For example, female HART was observed in a
mounted pair three times over a period of five days with male ZEB in September 2008 (Table
4.2); she had no clutch or other mating observations in 2009, but ZEB sired her entire clutch in
2010 (Table 4.5). Another male, BIPA, sired all of VLMAs offspring in 2009 and one offspring in
VLMAs multiply sired clutch in 2010 suggesting that the one offspring in 2010 clutch was
fertilized by stored sperm from the previous mating event. Further, repeat paternity between
years was observed by three additional males (DRLD, EZRA, and JAY) and occurred in ten of 12
(83%) between-year, paired clutches. Only two females with singly sired paired clutches
switched sires between years.
Levels of gene flow

Inter-population mating attempts were observed on three different occasions: one male
(EZRA) from the Will 1 population was found mounting a female (LIMA) from the Will 2
population in 2008 and a Will 2 male (BIPA) was found mounting a Will 1 female (MAUD) in
2008 and 2009. However, one multiply sired clutch (26-08) was the only genetic evidence of
male-mediated gene flow between populations.
139
DISCUSSION
In my study, behavioral observations were used to corroborate mating success and
variation in reproductive success. This information provided insight into the social mating
system as well as cryptic reproductive strategies such as mate choice and sperm storage. Using
complementary methods of field observations of mating behavior with genetic parentage analysis
of offspring, I was able to assess the mating system and reproductive success of E. blandingii.
During four consecutive years of radio-telemetry monitoring, I observed male and female E.
blandingii engaged in courtship behaviors with multiple individuals but parentage of clutches
collected during 2007-2010 was strongly skewed towards one male and multiple paternity was
rare. Male and females mated successfully with multiple individuals but successful matings did
not always correspond with previously observed mating attempts. In males, number of mates was
positively correlated with total number of offspring sired but I failed to detect inbreeding
avoidance in observed mating pairs or a decrease in hatching success in related pairs. Previous
chelonian mating system studies were important in documenting multiple paternity but were
often limited to one breeding season without behavioral data and could only infer number of
sires or assign paternity to candidate males (Valenzuela 2000, Ireland et al. 2003, Fantin et al.
2008, Refsnider 2009, Fantin et al. 2010). In such studies, it was not clear if multiply sired
clutches were a result of polyandrous mating system or instead, a mix of a seasonally
monogamous mating system with use of stored sperm from previous matings (Uller and Olsson
2008). More recent studies have examined parentage over multiple breeding seasons and have
included field data to test hypotheses regarding sperm storage and multiple paternity (Pearse et
al. 2001, 2002, Moore et al. 2009, Olsson et al. 2010, McGuire 2011, McGuire et al. 2011).
140
Timing and frequency of mating

Mating attempts in radio-telemetered E. blandingii were documented in every month
except June (nesting season) and during the over-wintering period (mid-November to midMarch). These findings are similar to anecdotal observations in previous E. blandingii studies
(Carr 1952, Graham and Doyle 1979). I observed most mating observations prior to
overwintering in the fall and during post-emergence in the spring. Fall mating observations have
been noted in Nova Scotia populations of E. blandingii (McNeil 2002) and could be attributed to
aggregations of turtles at overwintering sites (Newton and Herman 2009). A high occurrence of
mating has been documented in the spring for E. blandingii populations in Minnesota (Sajwaj
and Lang 2000) and the Great Lakes Region (Harding 1997).
I recorded the highest number of mating observations in July; however seven of the ten
July observations occurred in 2007. Most of the July 2007 courtship observations were clustered
in the same wetland area and involved three and four different males and females, respectively.
There was a white-tailed deer (Odocoileus virginianus) carcass in the marsh during that time that
seemed to attract several turtles to this particular location, presumably to scavenge. The high
density of turtles probably resulted in an increased encounter rate among individuals and the
observed flurry of mating activity. It appears that mating attempts between individuals occur
anytime there is an encounter and encounters are probably most frequent when turtles cluster or
aggregate within the same areas such as those used for overwintering.
Number of potential mates

Because males and females were both observed with multiple partners during radiotelemetry surveys, the social mating system of E. blandingii in Will County, Illinois appears to
141
be promiscuous. Further, I observed male E. blandingii more frequently with multiple mates than
females throughout the study. Similarly, Rovero et al. (1999) observed male European pond
turtles (Emys orbicularis) to mount multiple females and females were mounted by multiple
males. Kaufmann (1992) also reported that female wood turtles (Glyptemys insculpta) in central
Pennsylvania mated with multiple males between nesting seasons. In my study, some courtship
events were undetected. For example, there were 14 clutches sired by males with females for
which no courtship pairing was observed in the field. Although much effort went into radiotracking turtles multiple times a week, several mating events were likely undetected and
observed courtship displays probably only represent a subset of total mating attempts in the
population. Thus, number of courtship observations is likely to be underestimated and further
supports a promiscuous mating system classification.
Male reproductive success

Males and females successfully mated with multiple individuals during the course of this
study. McGuire (2011) and Refsnider (2009) also documented multiple mates for E. blandingii at
the E. S. George Reserve in Michigan and a Minnesota population, respectively. In my study,
although a total of nine males contributed offspring and two males sired offspring with at least
five different females, parentage was strongly skewed towards one male that sired 37% of all
offspring and 36% of all clutches. Disproportionate numbers of mates and skewed reproductive
success among males have also been documented in other reptile populations (Roques et al.
2006, Moore et al. 2009). For example in a population of E. orbicularis, one male sired 57% of
clutches (Roques et al. 2006). Although complete reproductive histories were missing for some
turtles, it is still apparent that reproductive success was not equal among male E. blandingii.
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Quantification of mating success

Conservatively, 41% of male mating attempts observed in the field resulted in sired
offspring. This success rate is lower than that of 84% observed in a population of tuatara
(Sphenodon punctatus), a seasonally monogamous reptile (Moore et al. 2009). In my study, over
half of courtship observations in the field were not successful either because 1) females were not
receptive and mounting behavior did not result in successful copulation, 2) post-copulation
cryptic female mate-choice prevented successful fertilization, or 3) the prevalence of variation in
sperm quality or sperm competition among males. In all observed courtship events, males
displayed mounting behavior but copulation success was uncertain based on observations alone.
In a group of wild-caught E. blandingii that were held in outdoor enclosures, only five
copulations were observed out of more than 100 mating attempts (Baker and Gillingham 1983).
Thus, low copulation frequency could explain why only a portion observed courtship events
resulted in successful matings in my study.

Until recently, few reptile studies have evaluated the relationship between male traits and
reproductive success (but see Kaufmann 1992, Pearse et al. 2002, Moore et al. 2009, Olsson et
al. 2010, Tuberville et al. 2011). I predicted that males with larger home range sizes would
encounter more females and subsequently have greater reproductive success. Only number of
mates was positively correlated with number of offspring. There was no relationship between
body size or home range size and reproductive success. Carapace length was also not a predictor
of reproductive success in C. picta (Pearse et al. 2002) but body size was a predictor of
reproductive success in S. punctatus (Moore et al. 2009), sand lizards (Lacerta agilis, Olsson et
143
al. 2010) and gopher tortoises (Gopherus polyphemus; Tuberville et al. 2011). Additionally,
dominance in G. insculpta, as measured by observations of agonistic interactions during malemale encounters, was positively correlated to reproductive success (Kaufmann 1992, Galbraith
and Kaufmann unpublished data). Amount of experience or age can also have an effect on
reproductive success: older or more experienced individuals sire more offspring (Tuberville et al.
2011). Other than the two sub-adult males excluded from the study, I was unable to assess age of
the remaining adult turtles because methods used to age young turtles such as annuli growth
rings become indiscernible beyond sexual maturity (Sexton 1959b). Finally, small sample size
may have obscured some relationships between male traits and reproductive success.

Females may be reluctant to mate with particular males if they could detect a fitness
disadvantage in the pairing, such as inbreeding (Amos et al. 2001, Stow and Sunnucks 2004,
Miller et al. 2010). Levels of relatedness varied among male-female pairs but I found no
indication that relatedness affected mating success or hatching success. Similar findings have
been observed in the grand skink (Oligosoma grande), a promiscuous lizard from New Zealand
(Berry 2006). Berry (2006) found that skinks mated with partners of varying relatedness but no
effect of offspring survival was evident. However, inbreeding avoidance has been has been
documented in fragmented populations of Cunninghams skink (Egernia cunninghami; Stow and
Sunnucks 2004).
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Detection of multiple paternity

Multiple paternity has been documented in many chelonian species (for recent reviews
see, Uller and Olsson 2008, Refsnider 2009, Davy et al. 2011) and range from 4% (McTaggert
2000) to as high as 100% (Valenzuela 2000, Ireland et al. 2003, Fantin et al. 2008, Fantin et al.
2010). However, variation in sample sizes, proportions of clutches sampled, and number of loci
used to infer paternity can make comparisons among studies difficult (Davy et al. 2011). In Will
County, Illinois, I detected only 8% multiple paternity in 28 clutches; levels much lower than
reported in previous studies of E. blandingii. Refsnider (2009) detected 56-81% multiple
paternity in 16 clutches and McGuire (2011) detected on average 47% multiple paternity in 77
clutches. Discrepancies in levels of multiple paternity among populations have also been
documented in studies of painted turtle (Chrysemys picta, 4-30%; McTaggert 2000, Pearse et al.
2001, 2002) and green sea turtle (Chelonia mydas, 9-61%; Fitzsimmons 1998, Lee and Hays
2004) that analyzed 18 clutches. Other freshwater chelonian studies with comparable sample
sizes have also detected low estimates (~10%) of multiple paternity (Pearse et al. 2006, Roques
et al. 2006). The low levels of multiple paternity detected in my study could be attributed to
lower population density and a female biased sex ratio (Stephens and Sunderland 1999).
My estimate of multiple paternity is conservative because I was unable to locate some of
the natural nests and some clutches were partially or completely depredated, thus some instances
of multiple paternity may have been missed. Further, clutch size affects detection of multiple
paternity (Kichler et al. 1999, Pearse et al. 2002, McGuire 2011) but I was able to detect multiple
paternity in a clutch with as few as three hatchlings. Interestingly, one male (BIPA) contributed
offspring to all three multiply sired clutches. Roques et al. (2006) also found that the same male
contributed to both multiply sired clutches in their study of E. orbicularis and postulated that this
145
male may be of higher quality. Thus, BIPA may have higher sperm quality compared to other
males, but little is known about sperm competition in turtles.
Hypotheses for the evolutionary advantages of multiple paternity include indirect female
benefits such as increased genetic variation of offspring (Pearse et al. 2002, Pearse and Anderson
2009). However, Uller and Olsson (2008) argue that evidence for such benefits is lacking and
that multiple paternity is more likely maintained by direct male fitness benefits from mating with
multiple females, low female mating costs, and sperm competition. In his classic paper, Bateman
(1948) noted that males are subject to stronger sexual selection and therefore should for strive for
greater numbers of mates and exhibit more variable reproductive success than females. The
results of my study are consistent with this pattern; number of mates and variation in
reproductive success were higher among male than female E. blandingii. Though sample size
precluded me from testing hatching success and genetic variation between single and multiply
sired clutches, hatching success of both single and multiply sired clutches varied drastically (12100%).
Evidence of sperm storage

Potential for sperm storage is high when females have multiple mates (Devine 1984).
Discovery of oviductal sperm storage (Gist and Jones 1989, Gist and Congdon 1998) and the
occurrence of offspring produced via stored sperm (Galbraith 1993, Pearse et al. 2001, 2002)
have been documented in many chelonian species. I documented one confirmed instance of
across-season sperm storage in E. blandingii when a male sired offspring the year after his death
but also noted additional occurrences of potential sperm storage from repeat paternities and from
comparisons of field observations with inferred parentage. Potentially, five clutches could have
146
resulted in fertilization using stored sperm 1-2 years after the observation. However, many of
these females sired a clutch by another male during the interim, which raises questions about
temporal viability in stored sperm and the effects of positional priority of sperm in the oviducts
(see below).
Repeat paternities have been documented in multiple species including C. picta (Pearse et
al. 2001, 2002, McGuire et al 2011), E. orbicularis (Roques et al. 2006), and E. blandingii
(McGuire 2011). In species that produce multiple clutches within the same nesting season, repeat
paternity is considered to be a result of sperm storage (Pearse et al. 2001, 2002, Roques et al.
2006, Sheridan 2010, McGuire et al. 2011) because of the low probability of a female remating
with the same male during the interval between oviposition of and ovulation of successive
clutches (Gist and Congdon 1998). Although stored sperm may also be used to fertilize clutches
among years (Pearse et al. 2001, 2002, Sheridan 2010, McGuire et al. 2011), there is greater
potential for remating to occur in the same pair of individuals between successive clutches for
species that only lay one clutch per year. For example, E. blandingii often use the same core
areas year after year (Congdon et al. 2011, Chapter One) and although turtles do not create pair
bonds, turtles that tend to use the same core areas likely encounter the same mates over time.
Thus repeated matings between some pairs of individuals are more likely than others.
Intraspecific territoriality or mate guarding could also result in high variation in reproductive
success and repeat matings (Emlen and Oring 1977). Some studies have documented
intraspecific aggressive behaviors or dominance hierarchies in male freshwater turtles (Kaufman
1992, Rovero et al. 1999) but strong evidence for territoriality is lacking. I observed five
instances of intrasexual mounting behavior between males during this study, involving five adult
147
males, one sub-adult male, and one juvenile male (unpublished data) but it is unclear whether
these observations were dominance displays or misdirected mating attempts.
It is also not clear if the three multiply sired clutches in this study were fertilized by
stored or recently inseminated sperm. The proportions of offspring sired by each male in the
multiply sired clutches varied within each clutch. Proportions of offspring sired by males should
depend on the contribution of sperm quantity or sperm quality of each male (Devine 1984).
There is conflicting evidence regarding the relationship of fitness effects and the use of stored
sperm. In E. orbicularis, use of stored sperm resulted in lowered hatching success and smaller
hatchling size (Roques et al. 2006). Conversely, use of stored sperm had no effect on hatching
success in C. picta (Pearse et al. 2002). In addition, the proportions of offspring sired by male C.
picta in multiply sired clutches depended on mate order of the males: the last male to mate sired
more offspring (Pearse et al. 2002). One instance of multiple paternity in my study supports this
pattern of sperm storage; one male (BIPA) sired an entire females clutch (38-09) in 2009 but
only sired one of seven offspring in that females clutch (38-10) the following year. The
inference being that BIPA was the last mate in 2009, but may not have mated (sperm storage) or
was other than last in 2010.
Conservation Implications
Because I was able to assign all offspring to eight sampled males and only one unsampled male, and individuals had a tendency to be recaptured multiple times during the radiotelemetry project (Dreslik et al. 2011), I suspect that most individuals in our populations have
been captured and that the number of adult samples is in accordance with actual population sizes
for the preserves. Both locations consist of small remnant populations (~18 and ~ten adult
148
individuals for Will 1 and Will 2, respectively) with limited dispersal and gene flow occurring
between them as evidenced by radio-telemetry surveys (Chapter One) and the parentage analysis
conducted in this study. High variation in reproductive success and low levels of multiple
paternity in the Will County populations compared to other E. blandingii populations (Refsnider
2009 and McGuire 2011) may be attributed to small population size, female biased sex ratios
(Stephens and Sunderland 1999), and disruption of the mating system (Lane et al. 2011).
Although males and females were observed with multiple partners, many of those mating
attempts were unsuccessful. Whether this was caused by female-mate choice, spermcompetition, or lack of mate encounters (i.e. Allee effects) is not clear. It appears that mating
success is related to number of mate encounters but identification of more refined determinants
of male reproductive success will require additional research and long-term monitoring. This
study examined reproductive success across four breeding seasons but this duration is still a short
window of time considering the longevity (70+ years) and reproductive potential of E.
blandingii.
Variation in reproductive success is thought to indirectly decrease effective population
size Ne (Nunny 1993, Anthony and Blumstein 2000) and can result in loss of genetic diversity
within just a few generations (Miller et al. 2009). Although I did not detect inbreeding avoidance
or lowered reproductive success in related pairs, skewed reproductive success among so few
individuals could have important genetic implications for the long-term persistence of remnant
populations (Frankham 1996) as well as management efforts such as captive rearing (Moore et
al. 2008b), reintroduction (Miller et al 2009), and translocation (Tuberville et al 2011). Further,
turtles have life history traits such as delayed maturity and low juvenile survival that exacerbate
declines and contribute to increased rates of genetic drift (Lee et al. 2011). Conservation plans
149
should seek to preserve or increase genetic variation for remnant turtle populations.
Unfortunately, the most feasible way to achieve this goal at these sites is captive breeding or
translocation of head-started individuals.
150
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160
TABLES
161
Table 4.1 Turtle ID, site (Will 1 or Will 2), start of tracking duration, end of tracking duration,
number of radio-locations (#Loc), carapace length in mm (CL) and 95% fixed kernel density
isopleth in ha (95K) for 29 E. blandingii radio-tracked at two preserves in Will County, Illinois
from 2006-2010.
ID
Site
Start
End
Females
EZMR
PRMA
JUDI
FRAN
BV
CLET
HART
MRTH
ETHL
EDNA
MILD
CLRA
MAUD
LIMA
BIMA
VLMA
SMLY
NOEL
HOPE
W1
W1
W1
W1
W1
W1
W1
W1
W1
W1
W1
W1
W1
W2
W2
W2
W2
W2
W2
28
14
23
15
2
12
12
14
2
6
18
14
16
6
11
23
24
17
22
MAY
JUN
JUN
JUL
JUL
JUL
JUL
SEP
MAY
JUN
JUN
NOV
MAY
MAY
MAY
APR
MAY
NOV
JUN
2006
2006
2006
2006
2006
2006
2006
2006
2007
2007
2007
2007
2008
2007
2007
2008
2008
2008
2009
11
30
30
2
23
8
27
30
23
7
30
30
30
20
30
30
15
15
30
JUN
MAY
MAY
MAY
NOV
AUG
MAY
MAY
MAY
JUN
MAY
MAY
MAY
NOV
MAY
MAY
APR
APR
MAY
Males
ZEB
MNGO
DRLD
VERN
JAY
EZRA
RMEO
AXEL
BIPA
LIPA
W1
W1
W1
W1
W1
W1
W1
W2
W2
W2
14
6
17
29
16
29
26
7
23
3
SEP
SEP
SEP
MAR
APR
JUL
AUG
MAY
JUL
AUG
2006
2006
2006
2007
2007
2007
2009
2007
2007
2007
19
15
13
11
19
21
13
14
14
21
OCT
OCT
OCT
SEP
OCT
OCT
OCT
OCT
OCT
OCT
# Loc
CL
95K
2008
2010
2010
2007
2009
2007
2010
2010
2010
2010
2010
2010
2010
2009
2010
2010
2010
2010
2010
277
475
435
113
463
175
293
419
359
155
293
234
219
375
362
236
208
78
42
195
200
212
199
200
230
209
214
217
210
194
211
208
195
205
188
201
207
205
9.5
18.4
22.8
10.4
20.7
13.2
11.6
8.8
8.4
10.2
13.1
16.2
9.3
12
11.9
10.1
20
9.6
11.8
2009
2009
2009
2007
2009
2009
2009
2009
2009
2009
394
368
260
104
302
276
10
266
309
287
221
233
218
226
234
221
184
185
230
196
19.4
25.1
19.3
12.6
23.7
19.1
.
8.7
15.6
14.6
162
Table 4.2 Mating attempts documented between 8 male and 15 female E. blandingii during
radio-telemetry from 2006-2009 in Will County, Illinois. The relatedness coefficient (R) for each
pair was calculated using ML-RELATE (Kalinowski et al. 2006). Mating attempts were
considered successful if the observed pair parented a clutch during the nesting season following
the observation. Mating success of observed pairs is denoted by a ? when mating success could
not be determined because nests could not be located, transmitters, or clutches had 100% hatch
failure. Mating success is denoted by N/A when females did not produce a clutch during the
nesting season following a documented mating attempt that occurred in the preceding fall or
spring. The potential for sperm storage was noted for pairs that had no apparent mating success
in the subsequent nesting season but that successfully produced clutches 1 year after the
observation.
Observed Pair
BIPA
BIPA
BIPA
BIPA
DRLD
EZRA
JAY
JAY
JAY
JAY
JAY
JAY
JAY
JAY
JAY
JAY
LIPA
LIPA
LIPA
LIPA
LIPA
LIPA
LIPA
MNGO
MNGO
MNGO
VERN
VERN
VERN
VERN
BIMA
MAUD
MAUD
VLMA
JUDI
LIMA
BV
BV
CLET
CLET
EZMR
FRAN
HART
JUDI
MRTH
CLRA
BIMA
BIMA
LIMA
NOEL
VLMA
VLMA
VLMA
EDNA
ETHL
MAUD
BV
CLET
EZMR
FRAN
Month
JULY
MAY
MAY
SEPT
APRIL
JULY
OCT
APRIL
JULY
AUG
JULY
APRIL
MAY
SEPT
JULY
NOV
OCT
OCT
OCT
NOV
SEPT
OCT
APRIL
SEPT
MAY
APRIL
JULY
APRIL
APRIL
MARCH
Year
Mating
Success
2007
2008
2009
2008
2008
2008
2007
2009
2007
2007
2007
2007
2008
2008
2007
2007
2007
2008
2007
2008
2008
2008
2009
2008
2007
2009
2007
2007
2007
2007
0.00
0.00
0.00
0.00
0.26
0.00
0.00
0.00
0.06
0.06
0.00
----0.00
0.00
0.16
0.11
0.00
0.00
0.13
0.16
0.00
0.00
0.00
0.00
0.00
0.00
0.00
0.09
0.06
-----
?
YES
?
YES
NO
?
NO
NO
?
?
NO
?
YES
YES
NO
YES
YES
?
YES
YES
NO
NO
NO
N/A
?
?
NO
?
YES
?
Potential
Sperm Storage
2 YRS
.
2 YRS
2 YRS
1 YR
2 YRS
163
Table 4.2 (Cont.)

Observed Pair
VERN
ZEB
ZEB
ZEB
ZEB
ZEB
ZEB
ZEB
ZEB
FRAN
BV
CLET
CLET
EZMR
HART
JUDI
MRTH
BV
Month
Year
Mating
Success
APRIL
OCT
JULY
JULY
SEPT
SEPT
MAY
JULY
SEPT
2007
2007
2007
2007
2006
2008
2009
2008
2006
----0.38
0.00
0.00
0.00
0.00
0.00
0.12
0.38
?
NO
?
?
NO
N/A
NO
NO
?
Potential
Sperm Storage
2 YRS
2 YRS
164
Table 4.3 Number of potential mates and number of mating attempts observed during radiotelemetry surveys conducted from 2006-2009 as well as number of successful mates, clutches,
and offspring inferred from parentage analysis from 2007-2010 for 19 female and 11 male E.
blandingii from two forest preserves (Will 1 and Will 2) in Will County, Illinois. Females FRAN
and CLET were radio-tracked for only a short duration and excluded from further analyses. Male
UNKN was an un-sampled male detected during paternity analysis.
# Potential # Mating
Mates
Attempts
Name
Site
Females
EZMR
MRTH
ETHL
EDNA
PRMA
MILD
CLRA
MAUD
JUDI
FRAN*
BV
CLET*
HART
BIMA
LIMA
VLMA
SMLY
NOEL
HOPE
W1
W1
W1
W1
W1
W1
W1
W1
W1
W1
W1
W1
W1
W2
W2
W2
W2
W2
W2
3
2
1
1
0
0
1
3
3
2
3
3
2
2
2
2
0
1
0
3
2
1
1
0
0
1
4
3
3
5
5
2
3
2
4
0
1
0
Males
MNGO
DRLD
VERN
JAY
EZRA
RMEO
ZEB
AXEL
BIPA
LIPA
UNKN*
W1
W1
W1
W1
W1
W1
W1
W2
W2
W2
W2
3
1
4
8
1
0
6
0
3
4
.
3
1
5
10
1
0
8
0
4
7
.
# Successful
Mates
# Clutches
# Offspring
2
2
1
0
1
1
1
2
1
0
1
0
2
1
1
2
2
1
1
2
3
1
0
3
3
2
2
3
0
2
0
2
1
2
2
1
1
1
14
30
18
0
26
28
20
11
31
0
9
0
18
4
20
15
10
9
9
1
1
2
5
2
0
2
0
3
5
1
1
2
2
8
3
0
1
0
3
4
1
18
9
16
102
37
0
21
0
14
46
9
165
Table 4.4 Mean estimates of A) potential mating attempts and B) successful mating attempts for
19 female and ten male E. blandingii observed during radio-telemetry surveys conducted from
2006-2009 and inferred from parentage analysis of clutches obtained 2007-2010.
A)
Groups
Females
Males
Combined
2006 2007 2008 2009

0.1
0.9
0.8
0.3
0.7
2.0
1.7
0.7
0.3
1.3
1.0
0.4
All Years
1.5
3.8
2.2
Groups
Females
Males
Combined
2007 2008 2009 2010

0.8
0.9
0.9
1.0
0.4
1.5
1.6
1.3
0.5
1.1
1.1
1.1
All Years
1.3
2.6
1.7
B)
166
Incb.
Nat
Cap
Nat
Cap
Cap
Cap
Cap
Cap
Cap
Cap
Cap
Cap
Cap
Nat
Nat
Cap
Cap
Cap
Cap
Cap
Cap
Clutch
01-07
20-07#
27-07#
05-08
22-08
26-08*
02-08
03-08
24-08
11-08
07-08
01-08
16-08#
20-08
27-08+
05-09
22-09
26-09#
02-09
03-09
11-09
2007
2007
2007
2008
2008
2008
2008
2008
2008
2008
2008
2008
2008
2008
2008
2009
2009
2009
2009
2009
2009
Year
W1
W2
W2
W1
W1
W1
W1
W1
W1
W1
W1
W1
W1
W2
W2
W1
W1
W1
W1
W1
W1
Site
EZMR
LIMA
BIMA
BV
MILD
MAUD
PRMA
JUDI
CLRA
MRTH
HART
EZMR
ETHL
BIMA
LIMA
BV
MILD
MAUD
PRMA
JUDI
MRTH
VERN
--------DRLD
EZRA
BIPA/EZRA
JAY
JAY
JAY
VERN
JAY
ZEB
----LIPA
LIPA
DRLD
EZRA
----JAY
JAY
JAY
11
16
14
11
15
18
14
14
14
13
15
11
16
11
16
8
11
15
14
12
11
3
16
14
10
0
15
12
10
7
5
12
5
16
9
9
0
9
15
1
0
4
4
0
0
1
2
1
0
0
3
2
1
3
0
0
0
0
2
0
0
0
0
#Eggs #Unvbl. #Dead

4
0
0
0
13
2
2
4
4
6
2
3
0
4
7
8
0
0
13
12
7
#Live
8
0
0
1
16
3
2
4
7
8
3
6
0
4
7
8
2
0
13
12
7
0.06
--------0.09
0
0/0.01
0.52
0
0.11
0.07
0
0
----0
0
0.09
0
---0.52
0
0.07
167
0.36
0
0
0
0.87
0.11(0.11/0.0)
0.14
0.29
0.29
0.46
0.13
0.27
0
0.36
0.44
1
0
0
0.93
1
0.64
Hatch Success
Table 4.5 Incubation method (natural vs. captive nest), year of oviposition, site (Will 1 vs. Will 2), known mother (), putative sire(s)
(), number of eggs sampled (#Eggs), number of unviable eggs (#Unvbl.), number of dead/malformed hatchlings (#Dead), number of
live hatchlings (#Live), total number of hatchlings sampled (N), relatedness coefficient (R) for male-female pairs, and hatching
success for 35 clutches collected from 16 female E. blandingii in Will County, Illinois during 2007-2010. Clutches in bold indicate the
sire was observed with the female during the spring or fall preceding the clutch. An * indicates that a clutch was multiply sired. A
+ indicates that a clutch was partially depredated before complete emergence. A # indicates that a clutch had 100% hatch failure
and no sire could be determined.
Incb.
Cap
Cap
Cap
Nat
Nat
Nat
Cap
Cap
Cap
Cap
Cap
Cap
Cap
Cap
Clutch
24-09
16-09
43-09
38-09
41-09
40-09*
22-10
26-10
02-10
03-10
11-10
07-10
27-10
38-10*
Table 4.5 (Cont.)
2009
2009
2009
2009
2009
2009
2010
2010
2010
2010
2010
2010
2010
2010
Year
W1
W1
W2
W2
W2
W2
W1
W1
W1
W1
W1
W1
W2
W2
Site
CLRA
ETHL
HOPE
VLMA
NOEL
SMLY
MILD
MAUD
PRMA
JUDI
MRTH
HART
LIMA
VLMA
JAY
MNGO
UNKN
BIPA
LIPA
LIPA/BIPA
EZRA
EZRA
JAY
JAY
JAY
ZEB
LIPA
LIPA/BIPA
13
19
13
9
16
12
10
12
11
16
15
15
13
8
0
1
4
2
7
2
0
4
0
1
0
0
0
0
0
0
0
0
0
0
0
0
2
0
0
0
2
0
#Eggs #Unvbl. #Dead

13
18
9
7
9
10
10
8
9
15
15
15
11
8
#Live
13
18
9
7
9
10
10
8
11
15
15
15
13
8
N
0.11
0
---0
0.16
0/0.03
0
0.01
0.52
0
0.07
0
0.13
0/0
168
1
0.95
0.69
0.78
0.56
0.83(0.50/0.33)
1
0.67
0.82
0.94
1
1
0.85
1.00(0.88/0.13)
Hatch Success
Table 4.6 Number of alleles, observed heterozygosity (Ho), expected heterozygosity (He),
probability of violating Hardy-Weinberg equilibrium (P-HWE), and parentage exclusion
probability for 14 loci amplified for 33 adult E. blandingii sampled from two preserves in Will
County, Illinois. Significance level = 0.004 after Bonferroni correction.
Locus
Ho
He
P-HWE
Exclusion
Probability
BATC9
Eb09
Eb17
Eb19
GmuA18
GmuA19
GmuA32
GmuB08
GmuD121
GmuD21
GmuD55
GmuD70
GmuD87
GmuD93
12
9
6
3
2
5
2
3
6
2
5
11
8
2
0.818
0.758
0.727
0.667
0.212
0.688
0.182
0.152
0.455
0.121
0.727
0.906
0.667
0.333
0.837
0.729
0.675
0.617
0.351
0.755
0.165
0.169
0.592
0.213
0.670
0.804
0.618
0.351
0.17
0.84
0.35
0.42
0.03
0.35
1.00
0.22
0.02
0.05
0.47
0.50
0.76
0.65
0.68
0.53
0.42
0.32
0.15
0.52
0.08
0.09
0.38
0.10
0.43
0.64
0.35
0.15
169
FIGURES
170
Fig. 4.1 Total number of unique mounting behavior (i.e. potential mating) observations of malefemale pairs among eight male and 15 female E. blandingii during radio-telemetry surveys from
2006-2009 at two forest preserves in Will County, Illinois.
Potential Mating Observations
# Observations
12
10
8
6
4
2
0
Month
171
Fig. 4.2 Total number offspring sired by nine male E. blandingii from 2007-2010 at two forest
preserves in Will County, Illinois.
Number of Offspring Sired by Males

120
# Offspring
100
80
60
40
20
0
Turtle
172
Fig. 4.3 Total number clutches sired by nine male E. blandingii from 2007-2010 at two forest
preserves in Will County, Illinois.
Number of Clutches Sired by Males

12
# Clutches
10
8
6
4
2
0
Turtle
173
CHAPTER 5
SUMMARY
Landscape alteration has been identified as the primary cause of species declines and loss
of biodiversity worldwide (Ballie et al. 2004, Stuart et al. 2004, Thomas et al. 2004). The most
serious impacts of landscape alteration are those attributed to habitat fragmentation such as
habitat loss, decreased patch size, and isolation (Fahrig 2003, Ewers and Didham 2005). Habitat
fragmentation can have serious implications for the population viability of long-lived organisms
such as turtles (Gibbons et al. 2000, Mitchell and Klemens 2000, Beaudry et al. 2008) that have
life history traits, such as delayed maturation, that tend to exacerbate declines (Congdon and van
Loben Sels 1993, Congdon et al. 1993, 1994). In addition, variation in traits among turtle species
such as dispersal ability, body size, abundance, and ecological specialization are proposed to
influence response to fragmentation (Ewers and Didham 2005). Assessing such traits among
species will aid in understanding species-specific sensitivity to landscape change and assist in
conservation strategies for turtles.
The main goal of this study was to examine attributes of a turtle assemblage that could
create variation in connectivity and long-term persistence among species within a fragmented
landscape in northeastern Illinois. I studied five freshwater turtle species that occur in remnant
preserves within Lower Des Plaines River Valley (LDPRV). These species vary in traits (e.g.
longevity, rarity) that should affect how they are responding to recent (past 150 years)
anthropogenic fragmentation. I used contemporary techniques such as radio-telemetry as well as
biochemical tools such as nuclear markers to address the studys objectives.First, I analyzed
movement among Blandings turtle (Emydoidea blandingii) across three fragmented sites,
spotted turtle (Clemmys guttata) across two fragmented sites, and eastern musk turtle
174
(Sternotherus odoratus), common snapping turtle (Chelydra serpentina), and painted turtle
(Chrysemys picta) at one isolated site. I then examined habitat partitioning and measures of niche
breadth among those five species at the isolated site. Next, I assessed the genetic diversity and
gene flow among E. blandingii, C. picta, and C. serpentina among three fragmented and one
intact site within the LDPRV. Finally, I examined the mating system and reproductive success of
E. blandingii across two adjacent remnant sites.
Movement and home range estimates were larger for adult than juvenile E. blandingii. Of
the five species, differences between males and females were evident only in C. guttata, with
females having larger mean daily distances (MDD) and a greater number of core home range
areas (#C) than males. Such differences have been attributed to nesting forays of gravid females
in previous studies of C. guttata (Wilson 1994, Litzgus and Mousseau, 2004). In addition,
significant differences in MDD, minimum convex polygon home range estimates (MCP), and
home range length (HRL) were detected between sites in E. blandingii and C. guttata. Variation
in amount, type, and distribution of wetland area across sites likely explains why individuals
moved farther at one site versus another. I also detected differences in movement and home
range among species at the Will 3 site. Adult E. blandingii and C. picta had larger home range
estimates than C. guttata. In addition, all species moved greater daily distances (MDD) than C.
guttata. Studies have shown that E. blandingii are capable of making long overland forays (> 1
km) between wetlands and to nesting sites (Ernst and Lovich, 2009, Chapter One), whereas the
other species were more restricted to movements within wetlands. I observed three E. blandingii
to move between sites (Will 1 and Will 2) during this study. Although, E. blandingii are
considerably more vagile than the other species in this study, S. odoratus, C. serpentina, and C.
picta were also capable of making long distance aquatic movements ( 1 km) via the Des Plaines
175
River. Conversely, C. guttata made the shortest movements and had the lowest home range
estimates compared to all other species.
Patterns of macro- and micro-habitat use also varied among the five species. All species
used multiple macro-habitat types but the rare turtle species, E. blandingii and C. guttata, most
frequently used cattail marsh macro-habitats whereas the common species (C. picta, C.
serpentina, and S. odoratus) most frequently used pond macro-habitats. In addition, use of mesic
prairie, sedge meadow, river, and pond macro-habitats differed between C. guttata and common
species while only use of pond macro-habitats differed between E. blandingii and two common
species, S. odoratus, and C. picta. I found that species most strongly partitioned micro-habitat
along an axis comprised of water depth, water and vegetative surface cover, vegetation height,
and understory canopy cover.Proportion of organic substrates at radio-locations was also an
important variable that differentiated habitat use among species in this study; use of organic
substrates was highest among C. guttata and E. blandingii.In both levels of habitat analyses, C.
guttata appeared to be the most restricted in use of habitat and appears to be a habitat specialist
whereas E. blandingii and C. serpentina broadly and similarly used macro- and micro-habitats
and maintained a relatively large measure of niche breadth suggesting that they are habitat
generalists.My results suggest that C. guttata is most vulnerable to degradation of high quality
interior shallow cattail marsh, sedge meadow, and mesic dolomite prairie from siltation caused
by flooding of the Des Plaines River and that broad variation in water and vegetation microhabitat characteristics is necessary to support a diverse freshwater turtle community.
All species (E. blandingii, C. picta, C. serpentina) demonstrated moderate to high levels
of genetic diversity and no indication of inbreeding. In comparisons between the intact and
fragmented LDPRV sites, none of the species demonstrated lower levels of genetic diversity in a
176
fragmented site. Standardized comparisons of genetic divergence among species showed that E.
blandingii was more differentiated across sites than C. picta or C. serpentina and I only detected
significant pairwise FST differentiation in E. blandingii and C. serpentina. Although FST values
were low (0.018-0.029), E. blandingii were differentiated between the intact and each of
fragmented sites as well as between two of the fragmented sites. Conversely, significant
differentiation of C. serpentina between fragmented sites is likely a result of small sample sizes
across sites. Gene flow was male-biased in E. blandingii across the fragmented sites but patterns
of dispersal between males and females in C. picta and C. serpentina were not strong. I found no
evidence of genetic population bottlenecks in any species but simulations of future genetic
diversity suggest that E. blandingii is more vulnerable to loss of genetic diversity than C. picta or
C. serpentina.
During four consecutive years of radio-telemetry monitoring, I observed promiscuous
mating behavior in E. blandingii as both males and females engaged in mating attempts with
multiple individuals. A total of nine males contributed offspring. Two males sired offspring with
at least five different females but parentage was strongly skewed towards one male that sired
37% of all offspring and 36% of all clutches collected during 2007-2010. Male and females
mated successfully with multiple individuals but successful matings did not always correspond
with observed mating attempts; only 41% of male mating attempts observed in the field resulted
in sired offspring. In males, number of mates was positively correlated with total number of
offspring sired but I failed to detect a relationship between inbreeding avoidance in observed
mating pairs or a decrease in hatching success and relatedness between male and female pairs.
Potential for sperm storage is high when females have multiple mates (Devine 1984) and
sperm storage has been documented in freshwater turtle species (Galbraith 1993, Pearse and
177
Avise 2001, Pearse et al. 2001, 2002). Although I documented many repeat paternities in
clutches among years, I only documented one confirmed instance of across season sperm storage
in E. blandingii when a deceased male sired offspring the year after he died. I also detected only
8% multiple paternity in 28 clutches; levels much lower than reported in previous E. blandingii
studies (Refsnider 2009, McGuire 2011). High variation in reproductive success and low levels
of multiple paternity in the Will County populations compared to other E. blandingii populations
(Refsnider 2009 and McGuire 2011) may be attributed to small population size, female biased
sex ratios (Stephens and Sunderland 1999), and disruption of the mating system (Lane et al.
2011).
Conclusion
Characteristics examined in this study including vagility, niche breadth, genetic diversity,
and reproductive success are all elements that contribute to the viability of species in altered
landscapes (Ewers and Didham 2005). Each species in this study has its own unique combination
of traits and requirements that should affect how it is responding to the recent anthropogenic
habitat loss and fragmentation within the LDPRV. Variation in abundance (endangered vs.
common) and life history traits such as generation time and reproductive frequency vary among
these species and will affect population growth accordingly.
The response of state-listed species, E. blandingii and C. guttata, is of particular concern
to wildlife management agencies in this region. For E. blandingii, the ability for long distance
movements (Chapter One) and broad niche breadth, including the use of the Des Plaines River
(Chapter Two), should increase connectivity in the fragmented landscape of the LDPRV.
However, remnant sites have relatively few E. blandingii (Banning 2006, Banning et al. 2006,
178
Dreslik et al. 2011) and evidence for loss of genetic diversity and genetic drift (Chapter Three).
Thus, successful dispersal among sites is probably limited at best. Reproductive success in this
species was skewed but inbreeding avoidance and inbreeding effects were not apparent
suggesting that loss of genetic diversity may not be an immediate threat in such a long-lived
species. For C. guttata, restricted use of high quality marsh and sedge meadow habitats put it at
greater risk of habitat degradation (e.g. siltation) but its smaller home range size and shorter
movements compared to the other species may actually decrease its vulnerability to isolation
than the other species. Genetic diversity and differentiation has yet to be assessed for this
species.
The common species, C. picta, C. serpentina, and S. odoratus, appear to be capable of
long distance movements via the Des Plaines River (Chapter One) and generally use habitats of
less quality (Chapter Two) indicating that they are more resilient to habitat degradation and
isolation. However, unlike E. blandingii, no marked individuals of these species have been
recaptured at sites other than their site of original capture (Dreslik et al 2011). Although I found
no evidence of loss of genetic diversity in C. picta, genetic patterns across sites were less clear
for C. serpentina (Chapter Three). Additional sampling is needed to confirm subtle instances of
divergence that I observed in C. serpentina.
179
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Species: A global species assessment. IUCN, Gland, Switzerland and Cambridge, UK.
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Banning W.J. .2006. Ecology of the Blandings Turtle (Emydoidea blandingii) at a Northeastern
Illinois Wetland Community. Unpublished. Masters Thesis, University of Illinois Urbana
Champaign, Illinois.
Banning W.J., M.J. Dreslik, C.A. Phillips. 2006. Continued study of the ecology of the
freshwater turtle community at Lockport Prairie Nature Preserve: With special emphasis
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183
APPENDIX A
SPATIAL METRICS FOR EMYDOIDEA BLANDINGII
Spatial metrics for 80 E. blandingii radio-tracked at Will County, Illinois from 2005-2010.
Listed are: turtle identification number (#), site, sex, carapace length (CL), start of tracking
duration, end of tracking duration, number of radio-locations (#Loc), minimum convex polygon
(MCP), mean daily distance moved (MDD), 95% fixed kernel density isopleth (95K), 50% fixed
kernel density isopleth (50K), number of 50% fixed kernel density isopleths (#C), and home
range length (HRL).
#
Site
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
44
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
KPFP
Sex CL
(mm)
F
F
F
F
F
F
F
F
M
M
F
M
J
M
J
F
M
F
J
J
J
F
M
F
M
F
F
F
J
195
200
212
199
200
230
209
177
221
233
214
218
155
226
141
217
234
210
156
128
150
194
221
211
180
208
195
181
106
Start
28
14
23
15
2
12
12
7
14
6
14
17
27
29
23
2
16
6
25
7
7
18
29
14
26
16
23
10
5
May
Jun
Jun
Jul
Jul
Jul
Jul
Jul
Sep
Sep
Sep
Sep
Oct
Mar
Apr
May
Apr
Jun
May
Jul
Jul
Jun
Jul
Nov
Apr
May
Mar
Apr
Jul
2006
2006
2006
2006
2006
2006
2006
2006
2006
2006
2006
2006
2006
2007
2007
2007
2007
2007
2007
2007
2007
2007
2007
2007
2008
2008
2009
2009
2009
End
11
30
30
2
23
8
27
20
19
15
30
13
30
11
15
23
19
7
15
15
15
30
21
30
21
30
21
13
15
Jun
May
May
May
Nov
Aug
May
Jul
Oct
Oct
May
Oct
May
Sep
May
May
Oct
Jun
Oct
Oct
Oct
May
Oct
May
Oct
May
Jul
Oct
Oct
# Loc MCP MDD 95K 50K #C HRL

(ha) (m) (ha) (ha)
(m)
2008
2010
2010
2007
2009
2007
2010
2006
2009
2009
2010
2009
2010
2007
2007
2010
2009
2010
2009
2009
2009
2010
2009
2010
2009
2010
2009
2009
2009
277 13.5
475 26.0
435 49.6
113 14.2
463 29.7
175 30.2
293 43.2
10
.
394 37.6
368 94.2
419 23.6
260 40.9
419 62.4
104 14.4
19
.
359 20.9
302 73.4
155 34.2
334 13.1
328
7.1
145
2.7
293 33.2
276 109.3
234 21.5
230 26.4
219 10.4
60
8.4
80 24.9
27
.
30.8
29.7
48.4
16.1
29.3
49.9
31.9
.
43.0
79.4
31.0
56.6
35.4
50.7
.
39.3
71.9
59.0
37.3
27.9
28.0
41.1
94.4
38.2
47.5
33.7
44.7
43.9
.
9.5
18.4
22.8
10.4
20.7
13.2
11.6
.
19.4
25.1
8.8
19.3
11.9
12.6
.
8.4
23.7
10.2
6.2
4.1
4.8
13.1
19.1
16.2
8.4
9.3
13.6
18.6
.
1.4
1.5
3.0
1.0
1.1
1.3
1.1
.
2.1
1.7
1.3
1.1
1.8
2.1
.
1.1
1.0
1.0
1.3
1.1
1.3
1.3
2.3
1.1
1.2
1.1
2.4
2.4
.
1
2
2
1
1
1
1
.
2
2
1
1
1
2
.
1
1
1
1
1
1
1
1
1
1
1
3
2
.
721.8
1108.8
1535.2
614.6
1060.8
1052.1
1366.2
.
960.3
2603.0
873.1
1127.8
2066.0
1043.9
.
1333.9
2149.5
1955.8
1275.8
741.9
327.7
1272.0
4231.9
995.7
1210.4
948.7
1277.5
1041.3
.
184
Site
45
46
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
21
22
23
24
25
26
27
35
36
39
42
46
47
53
63
90
7
16
KPFP
KPFP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
RPNP
RPNP
Sex CL
(mm)
M
F
F
M
M
J
F
F
F
F
J
F
F
J
J
J
J
F
M
F
F
F
J
F
F
M
M
F
F
J
M
J
J
J
F
F
J
J
184
199
195
211
217
114
206
192
198
174
139
131
196
100
143
127
104
200
194
204
203
178
131
198
216
212
187
206
210
119
233
148
153
114
195
215
110
145
Start
26
14
21
16
26
8
11
11
17
15
15
18
26
27
27
7
27
6
25
12
23
25
22
22
20
20
22
10
1
23
1
7
6
21
29
11
23
2
Aug
Sep
Apr
May
May
May
May
May
May
May
May
May
Jun
May
Jun
Jun
May
Jun
Jun
Jun
Jun
Jun
Jun
Jun
Jun
Jun
Jun
May
Apr
Apr
May
May
May
May
May
Jul
Apr
May
2009
2009
2005
2005
2005
2005
2005
2005
2005
2005
2005
2005
2005
2005
2005
2005
2005
2005
2005
2005
2005
2005
2005
2005
2005
2005
2005
2006
2006
2006
2006
2006
2006
2006
2006
2005
2007
2007
End
13
14
4
7
1
6
17
12
28
21
28
20
28
6
6
8
22
28
26
28
28
28
6
14
28
6
9
9
28
5
22
8
8
14
1
23
6
29

(ha) (m) (ha) (ha)
(m)
Oct 2009
Apr 2010
Jul 2007
Nov 2006
Sep 2006
Nov 2006
May 2007
Jul 2006
Jun 2007
Jun 2007
Jul 2005
Jun 2005
Jun 2007
Nov 2006
Nov 2006
Nov 2006
Sep 2006
Jun 2007
Jun 2006
Jun 2007
Jun 2007
Jun 2007
Nov 2006
Jul 2005
Jun 2007
Nov 2006
Aug 2005
Jun 2007
Jun 2007
Oct 2006
Mar 2007
Nov 2006
Nov 2006
Nov 2006
Jul 2006
Jun 2006
Oct 2009
Sep 2008
10
10
269
200
178
244
282
92
291
251
26
13
264
132
198
215
125
293
140
277
107
137
242
7
288
235
25
125
165
115
85
115
112
107
20
106
372
97
.
.
52.9
26.9
37.1
2.9
43.9
14.7
45.7
35.0
.
.
4.5
1.2
7.1
7.1
6.1
17.7
37.2
7.2
28.5
39.2
5.4
.
12.7
6.3
.
9.3
36.2
8.7
9.3
0.7
29.7
1.4
.
0.8
20.3
15.9
.
.
27.5
11.0
34.7
9.1
24.1
50.4
41.5
18.0
.
.
12.4
13.8
13.9
13.6
13.8
22.2
28.2
9.4
12.1
48.5
11.4
.
26.6
10.2
.
32.6
73.7
24.1
37.6
14.7
32.4
8.4
.
4.4
16.5
37.7
.
.
9.1
7.5
14.5
4.7
25.6
13.9
22.2
8.1
.
.
3.6
3.2
10.4
7.6
7.8
10.8
10.4
6.7
7.8
22.8
5.9
.
13.2
9.0
.
13.7
23.7
9.9
9.6
3.3
4.7
4.7
.
4.2
9.7
15.4
.
.
1.0
1.0
2.4
1.0
3.8
1.1
1.9
1.1
.
.
1.1
1.0
1.8
1.0
1.6
2.3
1.0
1.2
1.0
2.0
1.1
.
2.4
1.3
.
3.4
2.5
1.1
1.2
1.0
1.0
1.2
.
1.3
2.0
2.4
.
.
1
1
3
1
3
1
2
1
1
1
2
1
2
2
2
1
1
3
1
3
1
.
2
2
1
2
1
1
1
1
2
1
.
.
1534.3
797.8
1208.0
326.0
1218.2
1255.3
1263.2
803.9
.
.
498.8
228.7
418.9
794.9
659.7
895.0
1234.6
554.5
1173.7
1086.9
427.9
.
761.9
382.2
.
589.9
1429.0
1215.7
583.8
191.7
1124.4
297.9
.
191.1
841.1
753.8
185
Site
20 RPNP
26 RPNP
27 RPNP
28 RPNP
33 RPNP
34 RPNP
38 RPNP
40 RPNP
41 RPNP
42 RPNP
43 RPNP
44 RPNP
45 RPNP
Average
S.E.
Sex CL
(mm)
F
J
F
M
M
M
F
F
F
J
F
J
F
195
137
205
185
230
196
188
201
207
86
205
112
210
Start
6
13
11
7
23
3
23
24
17
23
22
22
17
May
May
May
May
Jul
Aug
Apr
May
Nov
Apr
Jun
Jun
Jul
2007
2007
2007
2007
2007
2007
2008
2008
2008
2008
2009
2009
2009
End
20
24
30
14
14
21
30
15
15
14
30
13
6
Nov
Feb
May
Oct
Oct
Oct
May
Apr
Apr
Oct
May
Oct
Aug

(ha) (m) (ha) (ha)
(m)
2009 375 30.1
2009 264 21.9
2010 362 19.4
2009 266 53.5
2009 309 151.9
2009 287 21.9
2010 236 76.3
2010 208 38.7
2010 78 31.3
2009 66
5.6
2010 42
7.7
2009 51
4.5
2009
8
.
197.4 27.5
14.1
3.2
42.6
21.0
33.0
37.1
60.5
52.4
52.0
66.2
39.6
20.9
20.3
15.6
.
33.8
2.3
12.0
7.5
11.9
8.7
15.6
14.6
10.1
20.0
9.6
7.5
11.8
5.5
.
11.8
0.7
1.6 1
2.0 1
2.1 1
1.2 1
1.1 1
1.6 1
2.9 1
3.8 1
1.1 1
1.5 1
2.9 1
1.1 1
.
.
1.6 1.4
0.1 0.1
901.8
1254.4
619.5
1946.2
2645.4
941.3
2733.9
1591.9
1345.1
360.1
435.2
394.0
.
1084.2
82.4
186
APPENDIX B
SPATIAL METRICS FOR CLEMMYS GUTTATA
Spatial metrics for 36 C. guttata radio-tracked at Will County, Illinois from 2005-2008. Listed
are: turtle identification number (#), site, sex, carapace length (CL), start of tracking duration,
end of tracking duration, number of radio-locations (#Loc), minimum convex polygon (MCP),
mean daily distance moved (MDD), 95% fixed kernel density isopleth (95K), 50% fixed kernel
density isopleth (50K), number of 50% fixed kernel density isopleths (#C), and home range
length (HRL).
#
Site
19
20
34
37
38
43
44
45
48
49
50
52
96
1
2
3
4
5
6
8
9
10
11
12
13
14
15
17
18
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
LPNP
RPNP
RPNP
RPNP
RPNP
RPNP
RPNP
RPNP
RPNP
RPNP
RPNP
RPNP
RPNP
RPNP
RPNP
RPNP
RPNP
Sex CL
(mm)
F
M
F
F
F
M
F
M
M
F
M
M
F
F
M
M
M
F
F
M
F
F
M
F
F
M
F
M
M
108
137
112
107
108
102
108
113
107
103
103
102
108
98
89
103
95
86
96
101
106
100
108
104
96
89
105
100
101
Start
18
17
1
6
22
7
7
9
7
15
14
21
31
2
19
16
23
23
24
19
25
17
18
23
24
1
1
30
1
May
May
Jul
Apr
Apr
May
May
May
May
May
May
May
Jul
Apr
Mar
Apr
Apr
Apr
Apr
Mar
Apr
Apr
Mar
Apr
Apr
May
May
Apr
May
2005
2005
2005
2006
2006
2006
2006
2006
2006
2006
2006
2006
2006
2007
2007
2007
2007
2007
2007
2007
2007
2007
2007
2007
2007
2007
2007
2007
2007
End
14
30
14
14
16
14
6
14
31
14
14
14
5
18
12
6
13
17
14
21
12
15
4
14
1
16
14
10
30
Nov
Oct
Nov
Nov
May
Nov
Nov
Nov
Oct
Nov
Nov
Nov
Sep
Jun
Jun
Jun
May
Jun
Nov
Oct
Dec
May
Jun
Apr
May
Oct
Nov
Oct
Sep

(ha) (m) (ha) (ha)
(m)
2006
2005
2006
2006
2006
2006
2006
2006
2006
2006
2006
2006
2006
2008
2008
2008
2008
2007
2008
2008
2008
2008
2008
2008
2008
2008
2008
2008
2008
232
99
196
96
18
84
85
82
80
81
78
73
15
194
119
185
111
39
281
158
291
146
51
149
34
51
269
253
37
10.3
5.4
5.1
0.3
.
1.7
1.8
1.4
1.3
1.3
0.1
0.4
.
0.9
0.6
9.1
1.3
0.6
1.2
0.9
3.7
1.3
1.5
2.7
1.5
4.2
9.5
1.2
1.3
12.2
6.7
7.5
8.2
.
3.3
12.2
9.4
6.6
5.5
4.5
4.4
.
15.3
11.5
15.8
12.9
6.5
11.9
12.4
17.9
16.2
15.0
18.1
12.0
14.4
22.7
22.5
15.2
1.5
0.9
1.5
0.7
.
0.9
1.0
1.0
0.5
0.9
0.4
0.8
.
1.2
0.8
1.1
1.3
0.6
1.0
1.2
1.1
1.7
1.4
2.1
0.8
2.2
1.7
1.2
1.3
0.2
0.1
0.1
0.2
.
0.2
0.1
0.1
0.1
0.1
0.1
0.1
.
0.1
0.1
0.2
0.2
0.1
0.2
0.4
0.3
0.3
0.2
0.2
0.2
0.3
0.2
0.3
0.1
2
1
3
2
.
1
1
1
1
2
1
1
.
2
1
2
3
1
2
1
2
3
1
2
2
4
2
1
1
954.6
469.8
530.8
85.6
.
340.2
214.5
161.3
302.7
218.1
65.7
136.3
.
143.8
96.4
734.5
241.7
204.8
156.9
127.3
298.5
263.9
191.9
301.7
262.6
416.4
737.2
198.0
169.8
187
Site
19
24
29
31
35
36
37
RPNP
RPNP
RPNP
RPNP
RPNP
RPNP
RPNP
Average
S.E.
Sex CL
(mm)
M
F
M
F
M
M
F
96
105
104
95
83
96
103
Start
1
17
14
16
22
21
3
May
Apr
May
May
Aug
Sep
Oct
2007
2007
2007
2007
2007
2007
2007
End
21
29
20
14
13
11
8
Mar
Jul
Oct
Nov
Nov
Sep
Oct

(ha) (m) (ha) (ha)
(m)
2009
2008
2008
2008
2008
2008
2008
235
181
241
256
184
134
130
2.2
3.7
2.1
1.1
2.0
3.4
3.4
20.0
28.4
20.4
14.6
9.4
14.5
25.4
2.2
1.9
1.6
1.1
0.7
1.8
1.5
0.2
0.3
0.2
0.3
0.1
0.1
0.3
1
2
2
1
1
2
3
101.6
12.8
2.6
0.4
13.3
1.1
1.2
0.1
0.2 1.7
0.0 0.1
241.7
350.5
271.1
157.3
211.4
333.8
379.2
293.2
33.7
188
APPENDIX C
SPATIAL METRICS FOR STERNOTHERUS ODORATUS
Spatial metrics for 15 S. odoratus radio-tracked in Will County, Illinois from 2005-2006. Listed
are: turtle identification number (#), site, sex, carapace length (CL), start of tracking duration,
end of tracking duration, number of radio-locations (#Loc), minimum convex polygon (MCP),
mean daily distance moved (MDD), 95% fixed kernel density isopleth (95K), 50% fixed kernel
density isopleth (50K), number of 50% fixed kernel density isopleths (#C), and home range
length (HRL).
Site
28 LPNP
29 LPNP
30 LPNP
31 LPNP
32 LPNP
33 LPNP
55 LPNP
58 LPNP
64 LPNP
65 LPNP
71 LPNP
72 LPNP
75 LPNP
88 LPNP
89 LPNP
Average
S.E.
Sex CL
(mm)
M
F
M
M
F
F
M
F
F
F
M
F
F
M
M
93
101
97
102
114
111
122
119
109
122
107
111
109
101
105
Start
21
20
21
21
20
20
26
28
29
29
31
31
2
6
6
Jun
Jun
Jun
Jun
Jun
Jun
May
May
May
May
May
May
Jun
Jun
Jun
2005
2005
2005
2005
2005
2005
2006
2006
2006
2006
2006
2006
2006
2006
2006
End
21
10
24
22
22
6
10
25
26
26
26
25
26
16
25
Jun
Jul
Aug
Sep
Sep
Jul
Aug
Sep
Sep
Sep
Sep
Sep
Sep
Jul
Sep

(ha) (m) (ha) (ha)
(m)
2005
2005
2005
2005
2005
2005
2006
2006
2006
2006
2006
2006
2006
2006
2006
1
13
22
61
52
6
53
80
74
62
59
84
62
35
64
54.9
6.2
.
.
58.2
1.1
30.7
.
1.5
1.9
9.5
1.2
3.0
0.7
5.0
1.5
4.0
9.9
5.0
.
.
93.7
16.5
54.4
.
26.1
15.3
33.4
27.4
26.1
19.9
29.8
27.7
27.8
33.2
6.2
.
.
7.0
4.6
4.3
.
2.6
3.9
11.2
3.8
3.0
3.5
5.0
5.3
7.2
5.1
0.7
.
.
.
.
.
.
0.7 1 1970.8
1.5 2
205.7
0.8 1 1628.2
.
.
.
0.8 1
290.4
0.8 1
315.6
1.7 2
701.3
0.9 1
263.7
0.8 1
317.5
1.3 1
142.2
0.8 1
487.9
0.8 1
360.0
0.9 2
367.7
1.0 1.3
587.6
0.1 0.1 169.7
189
APPENDIX D
SPATIAL METRICS FOR CHELYDRA SERPENTINA
Spatial metrics for 11 C. serpentina radio-tracked in Will County, Illinois in 2006. Listed are:
turtle identification number (#), site, sex, carapace length (CL), start of tracking duration, end of
tracking duration, number of radio-locations (#Loc), minimum convex polygon (MCP), mean
daily distance moved (MDD), 95% fixed kernel density isopleth (95K), 50% fixed kernel density
isopleth (50K), number of 50% fixed kernel density isopleths (#C) and home range length
(HRL).
Site
54 LPNP
56 LPNP
57 LPNP
59 LPNP
60 LPNP
61 LPNP
62 LPNP
73 LPNP
74 LPNP
86 LPNP
87 LPNP
Average
S.E.
Sex CL
(mm)
M
F
M
M
F
M
M
F
F
F
M
315
234
214
271
293
238
266
252
253
233
289
Start
24
26
26
29
28
28
30
1
1
28
2
May
May
May
May
May
May
May
Jun
Jun
May
Jun
2006
2006
2006
2006
2006
2006
2006
2006
2006
2006
2006
End
15
25
2
26
26
25
26
25
19
25
6
Sep
Sep
Jun
Sep
Sep
Sep
Sep
Sep
Jun
Sep
Aug

(ha) (m) (ha) (ha)
(m)
2006
2006
2006
2006
2006
2006
2006
2006
2006
2006
2006
83
72
7
75
23
75
82
80
17
65
42
56.5
8.7
0.8
8.9
.
3.7
10.2
4.1
0.2
2.0
.
11.4
10.8
5.8
1.5
16.6
38.5
.
15.8
64.3
37.2
5.4
15.6
.
50.6
66.3
34.5
7.5
2.5
3.6
.
1.4
8.5
2.3
1.3
3.2
.
6.9
6.4
4.0
0.9
0.7 1
0.4 1
.
.
0.4 1
0.6 2
0.4 1
0.4 1
0.4 1
.
.
0.7 1
1.0 1
0.5 1.1
0.1 0.1
160.1
541.6
.
467.9
839.2
419.1
160.2
221.6
.
985.5
965.1
528.9
110.4
190
APPENDIX E
SPATIAL METRICS FOR CHRYSEMYS PICTA
Spatial metrics for nine C. picta radio-tracked in Will County, Illinois in 2006. Listed are: turtle
identification number (#), site, sex, carapace length (CL), start of tracking duration, end of
tracking duration, number of radio-locations (#Loc), minimum convex polygon (MCP), mean
daily distance moved (MDD), 95% fixed kernel density isopleth (95K), 50% fixed kernel density
isopleth (50K), number of 50% fixed kernel density isopleths (#C), and home range length
(HRL).
Site
76 LPNP
77 LPNP
78 LPNP
80 LPNP
81 LPNP
82 LPNP
83 LPNP
84 LPNP
85 LPNP
Average
S.E.
Sex CL
(mm)
F
M
F
F
M
M
M
M
F
165
144
158
139
153
145
132
131
128
Start
1
1
4
5
2
2
2
2
2
Jun
Jun
Jun
Jun
Jun
Jun
Jun
Jun
Jun
2006
2006
2006
2006
2006
2006
2006
2006
2006
End
20
26
25
29
18
26
25
15
28
Jun
Sep
Sep
Jul
Sep
Sep
Sep
Sep
Jul

(ha) (m) (ha) (ha)
(m)
2006
2006
2006
2006
2006
2006
2006
2006
2006
19
21
56
37
60
58
68
59
20
44.2
6.6
.
17.9
3.0
0.7
6.0
2.7
7.6
3.4
7.9
6.1
1.9
.
205.4
35.9
20.9
62.7
27.6
22.1
36.2
15.1
53.2
22.4
.
15.3
7.4
5.1
9.5
11.0
11.6
8.0
10.0
9.8
1.1
.
2.1
2.0
1.8
2.0
3.6
1.8
1.9
1.8
2.1
0.2
.
1
1
1
1
1
1
1
1
1
0
.
1716.6
462.8
224.9
549.8
224.5
488.3
337.1
1599.6
700.4
213.4
191
APPENDIX F
SAMPLE SIZES FOR HABITAT PARTITIONING ANALYSES
Number of locations and sample sizes included in statistical analyses for five turtle species radiolocated from May-September 2006 at a preserve in Will County, Illinois.
Species
Male
Female
Total
E. blandingii
# Locations
(N)
318
(5)
1041
(13)
1359
(18)
C. guttata
# Locations
(N)
334
(5)
356
(5)
690
(10)
C. picta
# Locations
(N)
189
(4)
62
(2)
251
(6)
C. serpentina
# Locations
(N)
278
(4)
187
(3)
465
(7)
S. odoratus
# Locations
(N)
162
(4)
294
(5)
456
(9)
192
APPENDIX G
HABITAT PARTITIONING POST-HOC STATISTICAL RESULTS
P-values for MANOVA macro-habitat and ANOVA micro-habitat post-hoc tests between 18 E.
blandingii (EMBL), 10 C. guttata (CLGU), six C. picta (CHPI), seven C. serpentina (CHSE),
and nine S. odoratus (STOD) at a preserve in Will County, Illinois from May-September 2006.
Variables are as follows: mesic prairie (MP), floodplain (FP), river (R), marsh (M), sedge
meadow (SM), pond (P), principal component 1 (PC1), principal component 2 (PC2), and
proportion of organic substrates (SUB). Significance levels were accepted at = 0.05.
MANOVA Post-Hoc
Species 1 Species 2
EMBL
EMBL
EMBL
EMBL
CLGU
CLGU
CLGU
CHPI
CHPI
CHSE
CLGU
CHPI
CHSE
STOD
CHPI
CHSE
STOD
CHSE
STOD
STOD
MP
FP
0.06
0.81
0.77
0.69
0.01*
0.01*
0.00*
1.00
1.00
1.00
1.00
1.00
0.81
1.00
1.00
0.64
0.98
0.92
1.00
1.00
R
1.00
0.63
0.08
0.06
0.40
0.05*
0.04*
1.00
1.00
1.00
ANOVA Post-Hoc
SM
PD
PC1
PC2
SUB
0.50
0.99
1.00
0.75
0.69
0.87
0.06
1.00
0.98
0.84
0.18
0.00*
1.00
0.54
0.11
0.12
0.01*
1.00
1.00
1.00
0.12
0.00*
0.01*
0.00*
0.00*
0.00*
0.00*
1.00
1.00
1.00
0.00*
1.00
0.31
0.00*
0.00*
0.00*
0.00*
0.03*
0.83
0.36
1.00
0.00*
0.21
0.03*
0.87
0.81
0.35
0.15
0.03*
1.00
0.02*
0.00*
0.16
0.00*
0.00*
0.00*
0.00*
0.73
1.00
0.68
193
APPENDIX H
MULTIPLEX PANELS
Multiplex panel (MP#), primer, fluorescent labeling dye, and annealing temperature (TA) for
PCR reaction conditions designed for amplification of E. blandingii, C. picta, and C. serpentina.
E. blandingii & C. picta
Panel
Locus
Dye
TA
MP1
MP1
MP1
MP1
BATC9
GmuD70
GmuD121
GmuA19
6-FAM
VIC
NED
PET
58/48
58/48
58/48
58/48
MP2
MP2
MP2
MP2
MP3
MP3
MP3
MP3
GmuA32
GmuA18
Eb17
GmuD87
6-FAM
VIC
NED
PET
57
57
57
57
MP5
MP5
MP5
Panel
Locus
Dye
TA
GmuB08
GmuD90
GmuD55
GmuD21
6-FAM
VIC
NED
PET
58
58
58
58
Eb09
Eb19
GmuD93
6-FAM
VIC
NED
55
55
55
Dye
TA
6-FAM
VIC
NED
NED
51
51
51
51
C. serpentina
Panel
Locus
Dye
TA
Panel
MP6
MP6
MP6
MteC1
MteC112
MteD111
VIC
NED
PET
57
57
57
MP7
MP7
MP7
MP7
Locus
MteB103
MteD106
MteD2
MteD9
194
APPENDIX I
GENETIC DIVERSITY INDICES
Standard genetic diversity indices for loci that successfully amplified in E. blandingii, C. picta,
C. serpentina samples collected within the Lower Des Plaines River Valley. Listed are number
of samples (N) genotyped, detected number of alleles (#A), size range of alleles in base pairs,
number of private alleles (PA), observed heterozygosity (Ho), expected heterozygosity (He),
inbreeding coefficient (FIS), and probability of rejecting Hardy-Weinberg equilibrium (PHWE).
Bonferroni adjusted =0.004.
Locus
Size
AR
PAR
Ho
He
FIS
PHWE
3
5
9
11
5
3
2
5
2
2
6
10
4
2
98-107
94-112
137-167
146-184
140-160
194-200
139-145
145-165
119-121
198-230
196-248
225-265
177-193
153-157
3.0
4.0
6.6
8.1
4.8
2.3
1.8
4.9
2.0
2.0
4.5
8.5
4.0
2.0
0.00
0.50
1.34
1.57
0.07
0.01
0.00
0.01
0.00
0.00
0.83
1.58
0.00
0.00
0.727
0.727
0.773
0.818
0.500
0.136
0.091
0.714
0.136
0.318
0.636
1.000
0.773
0.136
0.599
0.632
0.724
0.818
0.663
0.129
0.087
0.745
0.325
0.325
0.618
0.821
0.687
0.268
-0.214
-0.150
-0.067
0.000
0.246
-0.056
-0.048
0.041
0.581
0.022
-0.030
-0.218
-0.125
0.490
0.323
0.225
0.962
0.353
0.044
1.000
1.000
0.038
0.016
1.000
0.880
0.209
0.584
0.058
3
4
8
7
5
3
2
4
2
2
5
6
5
2
98-107
94-109
129-163
146-182
140-156
197-203
139-145
145-155
119-121
198-230
196-252
217-257
177-213
153-157
3.0
4.0
8.0
7.0
5.0
3.0
2.0
4.0
2.0
2.0
5.0
6.0
5.0
2.0
0.00
0.01
1.31
0.51
0.01
1.01
0.01
0.08
0.00
0.00
1.26
0.17
1.00
0.00
0.727
0.727
0.818
0.636
0.636
0.182
0.455
0.727
0.455
0.182
0.818
0.727
0.455
0.182
0.579
0.640
0.769
0.727
0.632
0.244
0.351
0.698
0.483
0.165
0.657
0.736
0.508
0.298
-0.257
-0.135
-0.065
0.125
-0.007
0.254
-0.294
-0.041
0.060
-0.100
-0.245
0.011
0.106
0.389
0.312
0.113
0.948
0.260
0.254
0.143
1.000
0.518
1.000
1.000
0.395
0.930
0.225
0.277
Will 1 (E. blandingii)

Eb19
Eb17
Eb09
BATC9
GmuD121
GmuB08
GmuA32
GmuA19
GmuA18
GmuD93
GmuD87
GmuD70
GmuD55
GmuD21
22
22
22
22
22
22
22
21
22
22
22
21
22
22

Eb19
Eb17
Eb09
BATC9
GmuD121
GmuB08
GmuA32
GmuA19
GmuA18
GmuD93
GmuD87
GmuD70
GmuD55
GmuD21
11
11
11
11
11
11
11
11
11
11
11
11
11
11
195
Locus
Size
AR
PAR
Ho
He
FIS
PHWE
3
5
9
8
5
3
2
6
2
2
7
9
5
2
98-107
94-109
137-163
146-182
144-160
194-200
139-145
145-165
119-121
198-230
196-268
217-261
177-221
153-157
3.0
4.8
6.4
7.4
4.4
2.7
2.0
4.7
2.0
2.0
4.8
6.5
3.8
1.9
0.00
0.16
0.31
0.23
0.02
0.29
0.00
0.44
0.00
0.00
0.55
0.08
0.37
0.00
0.733
0.867
0.733
0.900
0.567
0.267
0.233
0.600
0.333
0.367
0.700
0.767
0.467
0.200
0.645
0.754
0.731
0.859
0.597
0.313
0.206
0.672
0.320
0.375
0.637
0.777
0.453
0.180
-0.137
-0.150
-0.004
-0.047
0.051
0.147
-0.132
0.107
-0.042
0.022
-0.099
0.013
-0.031
-0.111
0.867
0.728
0.489
0.791
0.809
0.258
1.000
0.137
1.000
1.000
0.301
0.498
0.538
1.000
3
5
10
8
7
3
2
8
3
2
7
13
5
2
98-107
94-109
129-163
146-186
140-164
194-200
139-145
145-169
119-123
198-230
196-240
217-269
177-217
153-157
2.5
4.8
6.0
7.3
5.2
2.3
1.6
5.9
2.2
1.9
5.4
9.3
4.0
2.0
0.00
0.02
0.17
0.73
0.45
0.01
0.00
1.44
0.22
0.00
1.70
0.90
0.40
0.00
0.490
0.633
0.551
0.796
0.653
0.265
0.082
0.714
0.265
0.163
0.714
0.776
0.653
0.286
0.510
0.733
0.644
0.852
0.609
0.235
0.078
0.766
0.262
0.150
0.732
0.875
0.550
0.273
0.040
0.137
0.144
0.066
-0.073
-0.130
-0.043
0.068
-0.012
-0.089
0.024
0.114
-0.187
-0.046
0.900
0.131
0.122
0.004
0.465
1.000
1.000
0.520
1.000
1.000
0.160
0.022
0.550
1.000

Eb19
Eb17
Eb09
BATC9
GmuD121
GmuB08
GmuA32
GmuA19
GmuA18
GmuD93
GmuD87
GmuD70
GmuD55
GmuD21
30
30
30
30
30
30
30
30
30
30
30
30
30
30
Grundy (E. blandingii)

Eb19
Eb17
Eb09
BATC9
GmuD121
GmuB08
GmuA32
GmuA19
GmuA18
GmuD93
GmuD87
GmuD70
GmuD55
GmuD21
47
47
47
47
47
47
47
47
47
47
47
47
47
47
196
Locus
Size
AR
PAR
Ho
He
FIS
PHWE
93
106
93
106
106
106
106
106
2
9
2
13
17
54
11
12
82-88
213-240
130-132
126-160
132-228
178-566
165-209
146-198
2.0
7.8
2.0
11.0
13.9
37.9
9.1
10.5
0.00
0.15
0.00
0.84
1.42
4.79
0.11
0.00
0.065
0.755
0.247
0.783
0.840
0.981
0.774
0.849
0.062
0.794
0.233
0.812
0.824
0.964
0.753
0.838
-0.033
0.049
-0.063
0.036
-0.019
-0.018
-0.028
-0.013
1.000
0.712
1.000
0.175
0.721
0.486
0.260
0.901
71
70
71
71
70
71
70
71
2
11
2
11
16
55
10
12
82-88
213-240
130-132
126-160
132-228
170-562
165-217
146-198
1.9
9.9
2.0
10.0
12.3
44.5
9.6
11.2
0.00
1.06
0.00
0.00
0.85
8.32
0.98
0.00
0.042
0.886
0.183
0.887
0.757
1.000
0.757
0.803
0.041
0.828
0.189
0.812
0.761
0.973
0.780
0.830
-0.022
-0.070
0.031
-0.093
0.005
-0.028
0.029
0.033
1.000
0.872
0.562
0.650
0.624
0.961
0.446
0.055
110
110
110
110
109
110
110
110
2
9
2
13
16
51
11
11
82-88
213-237
130-132
126-160
132-216
166-436
165-209
146-194
2.0
8.5
2.0
11.8
11.4
39.2
10.3
10.2
0.00
0.34
0.00
1.32
1.07
3.71
1.01
0.00
0.152
0.750
0.152
0.866
0.757
0.964
0.821
0.830
0.140
0.786
0.140
0.848
0.768
0.969
0.776
0.843
-0.082
0.046
-0.082
-0.021
0.015
0.005
-0.058
0.016
1.000
0.009
1.000
0.194
0.878
0.292
0.626
0.583
Will 1 (C. picta)

Eb17
GmuB08
GmuA32
GmuA19
GmuD93
GmuD70
GmuD55
GmuD21
Will 2 (C. picta)
Eb17
GmuB08
GmuA32
GmuA19
GmuD93
GmuD70
GmuD55
GmuD21
Will 3 (C. picta)

Eb17
GmuB08
GmuA32
GmuA19
GmuD93
GmuD70
GmuD55
GmuD21
197
Locus
Size
AR
PAR
Ho
He
FIS
PHWE
2
8
2
10
11
38
9
12
82-88
213-237
130-132
126-160
132-200
166-464
165-205
146-198
2.0
8.0
2.0
10.0
10.9
37.7
9.0
12.0
0.00
0.01
0.00
0.03
0.66
2.36
1.07
0.15
0.022
0.667
0.222
0.800
0.756
0.978
0.800
0.773
0.022
0.755
0.198
0.831
0.777
0.960
0.745
0.826
-0.011
0.117
-0.125
0.037
0.028
-0.018
-0.074
0.064
.
0.247
1.000
0.861
0.104
0.576
0.605
0.681
6
2
5
7
11
128-156
140-144
342-454
238-270
160-287
5.3
1.5
4.1
6.4
8.5
0.01
0.18
0.01
0.00
0.07
0.619
0.048
0.524
0.619
0.905
0.737
0.046
0.434
0.813
0.865
0.160
-0.024
-0.206
0.238
-0.046
0.095
.
1.000
0.118
0.621
6
2
6
9
13
128-156
140-144
342-454
238-278
160-287
5.8
1.6
5.0
6.3
8.5
0.08
0.02
0.20
0.29
0.46
0.886
0.086
0.743
0.829
0.914
0.809
0.082
0.720
0.793
0.852
-0.095
-0.045
-0.031
-0.045
-0.073
0.999
1.000
0.073
0.658
0.537
6
2
4
7
15
128-156
140-144
374-454
242-270
160-283
5.7
1.9
3.5
6.2
11.5
0.03
0.18
0.00
0.59
2.01
0.765
0.176
0.294
0.706
0.941
0.753
0.161
0.311
0.760
0.903
-0.016
-0.097
0.056
0.071
-0.042
0.873
1.000
0.411
0.953
0.953
Grundy (C. picta)

Eb17
GmuB08
GmuA32
GmuA19
GmuD93
GmuD70
GmuD55
GmuD21
44
44
44
44
44
44
44
43
Will 1 (C. serpentina)

MteB103
MteC1
MteC112
MteD9
MteD111
21
21
21
21
21

MteB103
MteC1
MteC112
MteD9
MteD111
35
35
35
35
35
.

MteB103
MteC1
MteC112
MteD9
MteD111
17
17
17
17
17
198
Locus
Size
AR
PAR
Ho
He
FIS
PHWE
5
1
5
8
10
128-153
140
342-454
238-270
160-287
5.0
1.0
5.0
8.0
10.0
0.01
0.00
0.31
0.55
0.38
0.700
0.000
0.700
0.700
1.000
0.770
0.000
0.635
0.815
0.875
0.091
.
-0.102
0.141
-0.143
0.420
.
0.137
0.215
0.800
Grundy (C. serpentina)

MteB103
MteC1
MteC112
MteD9
MteD111
10
10
10
10
10
199
APPENDIX J
NUMBER OF POTENTIAL AND SUCCESSFUL MATES
Number of potential mates observed during radio-telemetry surveys conducted from 2006-2009
and number of successful mates inferred from paternity analysis from clutches sampled from
2007-2010 in 19 female and 10 male E. blandingii in two preserves in Will County, Illinois. A
. indicates that a turtle was not tracked during a time period or was excluded from analyses.
# Potential Mates
Turtle
2006
2007
2008
2009
# Successful Mates
Total
2007
2008
2009
2010
Total
FEMALES
EZMR
1
2
0
.
3
1
1
.
.
2
MRTH
0
1
1
0
2
1
1
1
1
2
ETHL
.
1
0
0
1
1
1
1
1
1
EDNA
.
0
1
0
1
0
0
0
0
0
PRMA
0
0
0
0
0
1
1
1
1
1
MILD
.
0
0
0
0
.
1
1
1
1
CLRA
0
1
0
0
1
.
1
1
1
1
MAUD
.
.
1
2
3
.
1
1
2
2
JUDI
0
0
2
0
3
1
1
1
1
1
FRAN*
.
.
.
.
.
.
.
.
.
.
BV
0
3
0
1
3
0
1
1
.
1
CLET*
.
.
.
.
.
.
.
.
.
.
HART
0
0
2
0
2
.
1
0
1
2
BIMA
.
2
1
0
2
1
1
1
.
1
L IM A
.
1
1
0
2
1
1
1
1
1
VLMA
.
.
2
1
2
.
1
1
2
2
SMLY
.
.
0
0
0
.
1
2
.
2
.
.
1
0
1
.
.
1
.
1
NOEL
HOPE
.
.
.
0
0
.
.
1
0
1
-------------------------------------------------------------------------------------------------------------------------MALES
MNGO
0
1
1
1
3
0
0
1
0
1
DRLD
0
0
1
0
1
0
1
1
0
1
VERN
.
4
.
.
4
1
1
.
.
2
.
6
2
1
8
0
4
4
3
5
JAY
EZRA
.
0
1
0
1
0
2
1
2
2
RMEO
.
.
.
.
.
.
.
.
.
.
ZEB
2
2
2
1
6
1
1
0
1
2
.
.
.
.
.
.
.
.
.
.
AXEL
BIPA
.
1
2
1
3
0
1
2
1
3
L IP A
.
2
3
1
4
0
2
2
2
5
200

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SPATIAL ECOLOGY, HABITAT USE, GENETIC DIVERSITY, AND

REPRODUCTIVE SUCCESS: MEASURES OF CONNECTIVITY OF A

CHAPTER 1: Spatial ecology of a freshwater turtle assemblage in a fragmented landscape .....1

APPENDIX J: Number of potential and successful mates ..........................................................200

Estimation of spatial parameters

Within-species comparisons common species

Within-species comparisons common species

Sexton, O. J. 1995. Miscellaneous comments on the natural history of Blandings turtle

95% KDI (ha)

Fig. 1.1 (cont.)

Fig. 1.1 (cont.)

95% KDI (ha)

Fig. 1.2 (cont.)

50% KDI (ha)

Fig. 1.2 (cont.)

Fig. 1.3 (cont.)

Fig. 1.3 (cont.)

Fig. 1.4 (cont.)

= Proportional Similarity Index

= Proportion of radio-locations in macro-habitat i

= Proportion of available macro-habitat i

when habitat used is specialized.

= Percentage macro-habitat use overlap between species j and species k

species j and species k

To avoid correlation among micro-habitat variables, I conducted a principle components

= Ratio of the improvement of the optimal tree classification from chance

= # actual observations correctly classified by tree

Macro-habitat use.Qualitative assessments of wetland macro-habitat use vs. availability

Conservation Implications.Species-habitat relationships and dimensions of habitat

LITERATURE CITED (Formatted for the Journal Copeia)

Henle, K., D. B. Lindenmayer, C. R. Margules, D. A. Saunders, and C. Wissel. 2004.

Table 2.1 Proportions of available macro-habitat used by ten C. guttata (CLGU), 18 E.

Mean Habitat Rank

CLGU EMBL CHSE STOD CHPI

CLGU EMBL CHSE STOD CHPI

CLGU EMBL CHSE STOD CHPI

CLGU EMBL CHSE STOD CHPI

CLGU EMBL CHSE STOD CHPI

CLGU EMBL CHSE STOD CHPI

Water Depth (cm)

Vegetation Height (cm)

CLGU EMBL CHSE STOD CHPI

Fig. 2.3 (cont.)

DNA amplification, linkage disequilibrium, and Hardy-Weinberg equilibrium

Genetic diversity within species across sites

Genetic divergence within species

Genetic divergence among species

Future loss of genetic diversity

Genetic diversity within species across sites

Genetic divergence within species

Genetic divergence among species

Future loss of genetic diversity

Levels of genetic diversity

Measures of genetic divergence

Future loss of genetic diversity

Literature Cited (Formatted for the Journal of Molecular Ecology)

Gerlach G, Jueterbock A, Kraemer P, Deppermann J, Harmand P (2010) Calculations of

Kalinowski ST (2005) HP-Rare: A computer program for performing rarefaction on measures of

Parker PG, Whiteman HH (1993) Genetic diversity in fragmented populations of Clemmys

Dest (95% CIs)

GST (95% CIs)

My objectives were to determine 1) timing and frequency of mating attempts, 2) number

Determinants of male reproductive success

Genetic compatibility and reproductive success